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ASILIDAE 48 (Assassin Flies Or Robber Flies)

ASILIDAE 48 (Assassin Flies Or Robber Flies)

SURICATA 5 (2017) 1097

ASILIDAE 48 (Assassin or Robber Flies)

Jason G.H. Londt and Torsten Dikow

Fig. 48.1. Female of sp. with hymenopteran prey (Zambia) (photograph © R. Felix).

Diagnosis 164, 184), extending medially; face with mystax (Fig. 1), usu- ally macrosetose (Fig. 46), but sometimes only composed of Small- to very large-sized flies (body length: 4–65 mm; setae near lower facial margin (Fig. 200); positioned 1 length: 4–40 mm) (Figs 101, 185), that are predatory, capturing in dorsal ∕2 of head (Fig. 46); fore- and mid coxa positioned on the wing, and to a lesser extent, resting insects or close together; legs virtually originating at same level to cap- . ture and hold prey (Fig. 46); metakatepisternum usually small (Fig. 46), except in (Fig. 162), not visible between can be diagnosed as follows: labellum of proboscis mid and hind coxa. fused to prementum at least ventrally; hypopharynx heavily sclerotised, with dorsal seta-like spicules; labrum short, at most Head dichoptic in both sexes; face usually protruding to 1 ∕2 as long as labium; cibarium trapezoidal; vertex usually de- some extent, forming facial swelling (Fig. 1), but in several pressed (Figs 72, 73); postpronotal lobes fused to scutum (Figs taxa entirely plane (Fig. 200); face with mystacal macrosetae

Kirk-Spriggs, A.H. & Sinclair, B.J. (eds). 2017. Manual of Afrotropical Diptera. Volume 2. Nematocerous Diptera and lower . Suricata 5. South African National Institute, Pretoria; pp. 1097–1182. 1098 SURICATA 5 (2017) forming mystax (Fig. 1), which varies in extent from only cover- depressed (Figs 72, 80); all 3 ocelli circular, placed on single 1 ing lower facial margin (Fig. 200) to entire face (Fig. 46); frons ocellar triangle (Fig. 74); antenna positioned in dorsal ∕2 of predominantly parallel-sided (Fig. 80), but strongly divergent head (Fig. 46); antennal scapes separated proximally; postpedi- in a few taxa (Fig. 81); vertex slightly (Fig. 70) to considerably cel of varying length and shape (Figs 34, 46, 58, 66); stylus of

Figs 48.2–7. Photographs of living Asilidae (): (2) sp. ♂ (Zambia); (3) same ♀; (4) Apoclea inarticulata Theodor ♂ (United Arab Emirates); (5) Apoclea sp. ♀ (United Arab Emirates); (6) Congomochtherus penicillatus (Speiser) ♂ (Tanzania); (7) Dasophrys androclea (Walker) ♀ (South Africa). Figs 2, 3 (photographs © R. Felix), Figs 4, 5 (photographs © H. Roberts), Fig. 6 (photograph © S.A. Marshall), Fig. 7 (photograph © J.G.H. Londt).

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1099 varying development, shape and position, often composed of usually open (Figs 84, 145, 199), but often closed with stalk

1 or 2 cylindrical or rod-like articles, plus seta-like sensory ar- (Figs 83, 107, 233); cell m3 open (Fig. 199) or closed (Figs ticle distally (e.g., Asilinae, Fig. 1; Brachyrhopalinae, Fig. 18; 83, 84, 145); cell cua open (Fig. 199) or closed (Fig 83, 84); , Fig. 42; Ommatiinae, Fig. 46; , costal vein (C) usually circumambient (Figs 83, 84, 145, 199), Fig. 54; , Fig. 58; Tillobromatinae, Fig. 61; and sometimes terminating at veins CuP or CuA+CuP (Fig. 223)

Willistonininae, Fig. 66), or stylus reduced, only seta-like senso- and even further anteriorly at vein R4 (Fig. 221); auxiliary vein ry article present and positioned in cavity on postpedicel (e.g., usually absent (Fig. 199), sometimes appearing as short stump

Laphriinae, Fig. 33); proboscis well-developed and strong, usu- vein on vein R4 (Figs 85, 113, 223), in 4 genera long, connect- ally straight, extending beyond frontoclypeal suture (Fig. 46), ing veins R2+3 and R4 (Figs 84, 99, 201, 203); veins M1 and M2 single (Ancylorhynchus Berthold), with posteriorly curved never fused (Figs 83, 84, 199); alula usually well-developed proboscis resembling parrot’s beak (Fig. 110); labellum small, (Figs 83, 84, 123), reduced in size in several genera (Fig. 197) fused to prementum at least ventrally, usually entirely fused; and entirely reduced in a few genera (Fig. 171). palpus 1- or 2-segmented, usually small, cylindrical, sometimes laterally compressed and prominent, often setose. elongate, parallel-sided (Fig. 201), sometimes slightly tapering distally (Fig. 203), or constricted between Thorax with postpronotal lobe entirely fused to scutum (Figs tergites 2–3 or 2–4 (Figs 145, 199), or dorsoventrally flattened 164, 184), extending medially; cervical sclerite flat; proster- (Figs 86, 88, 222); tergite 2 usually wider than long (Figs 145, num dorsal margin with or without flange-like projection; pro­ 157, 199, 203), several genera with tergite 2 more than 5 × as sternum either fused to or separated laterally from proepister- long as wide (Fig. 172); abdominal tergites without anterodor- num; proepisternum fused to lateral postpronotum, setulose, sal apodemes; male and female tergites 1–8 well-developed; sometimes macrosetose; antepronotum usually bare, some- male and female sternites 1–8 usually well-developed and times setose and rarely macrosetose; postpronotum usually se- simple; tergites with (Figs 201, 227) or without lateral macro- tose, sometimes macrosetose; anterior proepimeron, anterior setae (Fig. 199); male terminalia usually straight (Fig. 194), but anepisternum, anterior and posterior basalare usually setose, often rotated up to 180° (Fig. 164); epandrium either separat- sometimes macrosetose (Fig. 1); posterior anepisternum, an­ ed and joined proximally (Figs 247, 278, 295), separated, not epimeron and anatergite setose or bare (Fig. 1); several taxa joined proximally (Fig. 285), or fused entirely medially (Figs with at least 1 macroseta on superoposterior angle of anepister- 265, 273); hypandrium usually well-developed and distinct num; katatergite setose or macrosetose, usually elevated and (Figs 240, 259, 278, 284, 292, 310), often reduced to varying convex; metakatepisternum usually small (Fig. 46), in Laphri­ degree (Fig. 271) or fused to gonocoxite forming gonocoxite inae, large and visible between mid- and hind coxa (Fig. 162); + hypandrial complex (Figs 268, 275, 281, 197); gonocoxite scutum usually smooth, sometimes punctate; dorsocentral se- of varying shape, often with -specific characters (Figs tae of varying length and development pre- and postsuturally 259, 262, 269, 287, 310); gonocoxal apodeme usually absent, (Figs 1, 61); acrostichal setae usually present presuturally; sev- sometimes present, either short or long; gonostylus of varying eral genera with scutal mane (Figs 10, 11, 194), composed of shape, often with species-specific characters (Figs 248, 261, dense setae between rows of acrostichal and dorsocentral se- 263, 269, 310); subepandrial sclerite usually without setae tae; noto­pleural, supra-alar and postalar setae usually present or protuberances; lateral ejaculatory apodeme present; 1–3 (Figs 1, 46, 58). Scutellum large; mediotergite (mesopostnotum) phallic prongs, usually short, sometimes long; female termi- not visible in dorsal view (Fig. 201); apical and discal scutel- nalia with tergite 8 usually simple, sometimes with anterior lar macrosetae absent (Fig. 200) or present (Figs 1, 142, 144). transverse apodeme (Fig. 306); usually consisting of Legs with hind coxa setose or macrosetose laterally (Fig. 1); segment 8 and following segments (Figs 23, 53), but sometimes hind coxa in a few taxa with anterior peg-like process; median involving segment 5 or 6 and following segments; ovipositor hind trochanter setose, sometimes expanded medially and for direct sand or soil oviposition, with tergite 10 divided into 2 macrosetose; femur usually cylindrical (Figs 1, 46) to at least acanthophorite plates with acanthophorite spines (Figs 22, 23, slightly expanded distally (Figs 22, 40), one genus with greatly 53, 56–58, 136, 306), a few genera with strong macrosetae on expanded fore femur (Fig. 52) and several genera with dis- for sand or soil oviposition, ovipositor for other types tinctly clubbed hind femur (Figs 34, 146, 228), sometimes with of oviposition, such as egg-dropping, simple (Figs 42, 45), for ventrodistal macrosetae mounted on tubercles; hind tibia usu- placing eggs in vegetation often knife-like in taxon-specific ar- ally straight, sometimes arched medially; proximal tarsomere rangements of sclerites (Figs 7, 17, 252, 254), for placing eggs as long as (Fig. 18) or sometimes longer than tarsomeres 2–3 in decaying wood, simple (Fig. 37); either 2 or 3 equally large, combined (Fig. 200); pulvillus usually well-developed (Fig. 1), poorly- to well-sclerotised spermathecae present, sometimes with two dorsal ridges, entirely reduced in several genera (Figs median spermatheca larger or smaller than lateral ones; genital 25, 45, 66), or less commonly to small lobe visible at base of fork usually either ring-like (joined anteriorly and posteriorly) claw; setiform empodium usually present, sometimes reduced or inverted U-shaped (joined anteriorly, separated posteriorly), in length and rarely laterally compressed. Wing membrane anterior and lateral genital fork apodemes usually present. usually hyaline (Fig. 195), sometimes darkened through brown staining (Figs 145, 159, 211), or patterned through white and (e.g., Figs 312–319) robust with 11 apparent segments, brown staining (Figs 123, 157); membrane with microtrichia with small, partly retractable head capsule (Figs 312, 313); head in many taxa, often covering entire or almost entire wing (Fig. dorso-ventrally flattened; maxilla dorso-ventrally flattened, di-

137) or with specific pattern; cell r1 open (Figs 189, 199) or vided into proximal part and movable distal part (Fig. 314); closed (Figs 83, 84, 145), when closed either by stalk (Fig. 145) maxillary palpus inserted sub-apically (Fig. 314); mandibles or R1 and R2+3 joining at point where they meet costal vein pointed, stylet-like, not extending to apex of maxilla (Fig. 314); (C) (Figs 83, 233); cell r4 open (Figs 83, 84, 145, 199); cell r5 labrum pointed; submentum forming ventral plate; abdominal

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1100 SURICATA 5 (2017)

Figs 48.8–13. Photographs of living Asilidae (Asilinae): (8) Eremisca heleni (Efflatoun) ♂ (left) and ♀ (right) in copula (United Arab Emirates); (9) Melouromyia natalensis (Ricardo) ♂ (South Africa); (10) Neolophonotus sp. ♂ (South Africa); (11) same; (12) N. abun­ tius (Walker) ♂ with heteropteran prey (South Africa); (13) Philodicus sp. ♂ with cicindelid prey (Namibia). Fig. 8 (photograph © H. Roberts), Fig. 9 (photograph © J.G.H. Londt), Figs 10, 11, 13 (photographs © S.A. Marshall), Fig. 12 (photograph © B. Londt).

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1101 segments 1–6 sometimes (Laphriinae) with contractile protu- non-predacious proboscis, as well as long antennae, peculiar berances dorsally and laterally and -like protuberances wing venation and resting position on the ground with the body ventrally (Fig. 313) otherwise smooth (Fig. 312); posterior spir- almost touching the surface are unlike most assassin flies. acle on apparent penultimate abdominal segment (Figs 312, 313); the caudal abdominal segment often appears partly di- vided and difficult to distinguish subdivision; apex of abdomen and immature stages with taxon-specific sclerotised (hook-like) appendages. An ecological classification has been attempted, based on (e.g., Figs 320–323) robust with appendages movable observations of Afrotropical species and the structure of their (exarate); head with 2 pairs of antennal processes; abdominal ovipositor (Londt 1994c). Most of the information available is segments 1–7 with row of posteroventrally-oriented processes that recorded on specimen labels and in taxonomic literature. dorsally; segment 9 with caudally-oriented terminal processes. However, an analysis of 26 of data from the semi-arid Karoo near Willowmore (Eastern Cape Province, South Africa) Asilidae are very distinct and should not be confused in the (Londt 1998a) and a -long survey undertaken in a single field with most . The three rarely collected species of mixed seasonal grassland habitat (KwaZulu-Natal Province, the closely related Apioceridae (see Chapter 46) superficially re- South Africa) (Londt 2002c) have been conducted. Dikow et semble ground-perching robber flies, but their non-predacious al. (2009: 327) compared the species richness and endemicity proboscis, short antennae and peculiar wing venation easily of Asilidae in biodiversity hotspots, wilderness areas and com- differentiate them from Asilidae. In addition, Afrotropical Api­ parable areas in the Afrotropical Region and concluded that ocera Westwood species run over the surface on slender legs the hotspots and wilderness areas sensu Conservation Inter- and do not sit or perch. Some Mydidae (see Chapter 47) could national (Mittermeier et al. 2003; Myers 2003), harbour more also be confused with certain Asilidae species, but, likewise, the Asilidae species than comparable areas outside.

Figs 48.14–17. Photographs of living Asilidae (Asilinae): (14) Promachus sp. ♀ with asilid prey ( sp. ♂) (Namibia); (15) same ♂ (left) and ♀ (right) in copula (Namibia); (16) same ♀ (South Africa); (17) Synolcus (Macquart) ♀ (South Africa). Figs 14, 15 (photographs © S.A. Marshall), Figs 16, 17 (photographs © J.G.H. Londt).

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1102 SURICATA 5 (2017)

Dennis et al. (2013) summarised known information on the usually being equipped with strong setae or spines with which biology and morphology of immature stages. Oviposition com- to dig), or eggs are deposited into suitable crevices in plant monly involves random egg-dropping, introduction of eggs di- material or pasted between leaves (females frequently being rectly into the ground (the of those species involved equipped with elongate, knife-like ovipositors). While studies

Figs 48.18–23. Photographs of living Asilidae (Brachyrhopalinae and ): (18) Afroholopogon capensis (Lindner) ♀ (South Africa); (19) Afroholopogon sp. (South Africa); (20) Habropogon bussinowi Bosák & Hradský ♂ (United Arab Emirates); (21) Pegesimallus aulicus (Wiedemann) ♀ (South Africa); (22) Pegesimallus sp., ovipositing in soil (South Africa); (23) same with isopteran prey (Zambia). Figs 18, 21 (photographs © J.G.H. Londt), Figs 19, 22 (photographs © S.A. Marshall), Fig. 20 (photo- graph © H. Roberts), Fig. 23 (photograph © R. Felix).

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1103 undertaken in other zoogeographical regions (e.g., Melin pupae. Some of the better known contributions are those of 1923; Musso 1978) reveal that larvae are predatory, feeding Engel (1929: larva and pupa of Hyperechia Schiner), Engel on subterranean invertebrates, very little detailed biological (1932: immatures of Proagonistes Loew), Engel & Cuthbertson information exists. Assassin larvae most probably live in soil (1934: immatures of Alcimus Loew and Hyperechia), Engel & or in burrows in wood, but knowledge of Afrotropical species Cuthbertson (1939: immatures of Neolophonotus Engel and is largely confined to isolated descriptions of eggs, larvae and Promachus Loew), Londt & Harris (1987: larvae of Millenarius

Figs 48.24–29. Photographs of living Asilidae ( and Laphriinae): (24) Oligopogon penicillatus Loew ♂ (South Africa); (25) Anypodetus sp. ♂ (Namibia); (26) A. fasciatus Hermann ♀ (Namibia); (27) sp. ♂ (South Africa); (28) Gonecca­ lypsis sp. ♂ (Tanzania); (29) Hoplistomerus nobilis Loew ♀ (South Africa). Figs 24, 26, 29 (photographs © T. Dikow), Figs 25, 27, 28 (photographs © S.A. Marshall).

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1104 SURICATA 5 (2017)

Figs 48.30–35. Photographs of living Asilidae (Laphriinae): (30) vorax Loew ♀ with buprestid prey (Oman); (31) Laphyctis sp. ♂ (Namibia); (32) Laxenecera albicincta (Loew) ♂ (left) and ♀ (right) in copula (South Africa); (33) Notiolaphria coerulescens (Macquart) ♀ (Madagascar); (34) same ♂ (Mauritius); (35) Nusa infumata (Loew) ♂ with lepidopteran prey (Tanzania). Fig. 30 (photograph © M. Borer), Fig. 31 (photograph © T. Dikow), Fig. 32 (photograph © J.G.H. Londt), Figs 33, 35 (photographs © S.A. Marshall), Fig. 34 (photograph © D. Martiré).

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1105 dichaetus (Hull, 1967) (Figs 318–319) and Londt (1991: eggs focusing on Asiloidea (Trautwein et al. 2010) and Diptera in and larvae of Damalis femoralis Ricardo, 1925). general (Wiegmann et al. 2011).

Adult flies are predators, feeding exclusively on living prey, Artigas & Papavero (1988), Geller-Grimm (2004), Hull (1962) most frequently caught in flight, but sometimes actively hunting and Papavero (1973) followed a traditional classification based resting prey (e.g., Leptogastrinae). The proboscis and internal on overall morphological similarity. Dikow (2009a) published sucking pump (cibarium) are well-developed for purpose the only phylogenetic classification of Asilidae to date, but did and harbour all of the morphological autapomorphies (see di- not provide an updated identification key to world subfamilies. agnosis, Dikow 2009a). Although there is scattered informa- While that study was unable to include all genera known at the tion concerning prey taken by assassin flies, Londt (1987a: 44, time (528), only 127 genera (24%) could not be placed in the 2006b) summarises what is known regarding adult feeding in phylogenetic classification, including 25 Afrotropical genera. general. of honeybees in particular (Londt 1993a) Since 2009, 13 Afrotropical genera have been described and and in general (Londt 1999c), have received at- along with the 25 previously unplaced genera are here placed tention, as well as the predation of robber flies by other robber within this most recent classification. No new phylogenetic flies (Fig. 14) (Londt 1995). In most instances, a broad range of analyses have been conducted, but genera are placed in one prey is taken (Figs 1, 13, 23, 30, 35, 42, 43, 54, 55, 58, 60), of the 11 subfamilies based on presence of synapomorphic but a few species appear to specialise (e.g., Hoplistomerus no­ character states. bilis Loew, 1858 (Fig. 29), primarily perches on large dung and feeds almost exclusively on small dung ). Of the 14 subfamilies recognised globally (Dikow 2009a), 11 are represented in the Afrotropical Region. Only the Australa- Male and female flies of the same species are morphologi- sian and Neotropical Bathypogoninae and Phellinae, as well as cally identical (except the structures pertaining to the termina- the Australasian and Holarctic Dioctriinae are not found with- lia) (Figs 187–190), with females sometimes slightly larger (Figs in the confines of the region. The most diverse subfamilies in 85–88). Only in very few Afrotropical species, e.g., Dasple­ terms of number of genera and species are the Asilinae (41 tis vespertilio (Engel, 1932) (Fig. 49) (Londt 2010a) and Lax­ genera, 574 species), Laphriinae (33, 201), Stenopogoninae enecera albicincta (Loew, 1852) (Fig. 32), is distinct sexual di- (22, 269) and Brachyrhopalinae (16, 101), while several sub- morphism evident. For the latter species, this dimorphism led families are only represented by a few genera, such as Dasy­ to the description of the male and female as different species, pogoninae (3 genera, 57 species), Stichopogoninae (6, 26), i.e., L. albicincta and L. zonata Loew, 1860, respectively. Males Tillobromatinae (2, 27) and Trigonomiminae (2, 61). of several Pegesimallus Loew species exhibit long, laterally compressed setae on the mid and hind tibiae and tarsi, giving The subfamily Asilinae is a megadiverse taxon in the Afro- them a “feathered” appearance, which led to the description tropics, comprising 41 genera (183 genera globally), 26 of of the genus Lagodias Loew and only later was it found to be which are endemic and 574 species. This subfamily is global an exclusively male, sexually dimorphous character (Oldroyd in distribution, except Antarctica. While most Afrotropical gen- 1974). Females and males of other species, e.g., Acnephalo­ era (29) include less than 10 species, Promachus (97 species) myia Londt species (Figs 85–88), exhibit some sexual dimor- and Neolophonotus (254) are extremely diverse and can be phism, but the sexes can be assigned to the correct species encountered throughout the region, with Neolophonotus be- despite such differences. ing particularly common in Southern Africa. Eleven endemic genera are monotypic and an additional nine have 2–4 species Mating takes place in taxon-specific positions and involves, each. Asilinae was newly delineated by Dikow (2009a), who e.g., standing -to-tail (Figs 15, 32, 55), perching tail-to-tail combined the and Asilinae sensu Artigas & Papave- (Fig. 63), or standing on top of each other (Fig. 59). ro (1988) and Geller-Grimm (2004) as Apocleinae was shown to be paraphyletic. The genus Loew still requires clar- ification in an Afrotropical context (see below). Economic significance The subfamily Brachyrhopalinae is a medium diverse taxon No species of Asilidae are known to have economic signif- in the Afrotropics, comprising 16 genera (currently 32 genera icance. Assassin flies, however, can have an effect on popu- globally), 13 of which are endemic and 101 species. The sub- lations of their prey insects and spiders, including honeybees family is distributed globally with the exception of the Ocean- (Londt 1993a). As predatory flies, Asilidae may be seen as in- ian Region and Antarctica. Thirteen Afrotropical genera have dicative of environmental health, i.e., greater numbers and di- less than ten species, three endemic genera are monotypic and versity of robber flies may indicate good environmental health. the endemic genera Afroholopogon Londt (19 species) and Rhabdogaster Loew (39) are relatively diverse. Brachyrhopal- inae was elevated to subfamily status by Dikow (2009a) and Classification includes genera previously assigned to Dasypogoninae and Stenopogoninae sensu Artigas & Papavero (1988) and Geller- Asilidae is placed in the superfamily Asiloidea and is closely Grimm (2004). related to Apioceridae and Mydidae. Dikow (2009a) sum- marised the history of placements of Asilidae within Asiloi- The subfamily Dasypogoninae is a very small taxon in the dea and proposed that a clade (Apioceridae + Mydidae) is Afrotropics, comprising three genera (17 genera globally), one sister-group to Asilidae, based on both morphological (Dikow of which is endemic and monotypic and 57 species. This sub- 2009a) and molecular data (Dikow 2009b). This set of rela- family is global in distribution, except Antarctica. ­ tionships has also been supported by molecular analyses inae was newly delineated by Dikow (2009a) as the monophyly

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1106 SURICATA 5 (2017)

Figs 48.36–41. Photographs of living Asilidae (Laphriinae): (36) Proagonistes praeceps (Walker) ♀ (South Africa); (37) Proagonistes sp. ♀ (Namibia); (38) Prytanomyia kochi (Lindner) ♀ (Namibia); (39) Stiphrolamyra sp. ♂ (United Arab Emirates); (40) Trichardis picta Hermann ♂ (Namibia); (41) T. leucocomus Wulp ♂ (United Arab Emirates). Fig. 36 (photograph © J.G.H. Londt), Fig. 37 (photograph © S.A. Marshall), Figs 38, 40 (photographs © T. Dikow), Figs 39, 41 (photograph © H. Roberts).

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1107 of the previous delineation sensu Geller-Grimm (2004) and all genera also occur outside the Afrotropical Region, primarily Papavero (1973) could not be established and several genera in the Old World, with Schildia Aldrich centred in the New were assigned to Brachyrhopalinae, none of which are Afro- World. tropical. The subfamily Ommatiinae is a diverse taxon in the Afro- The subfamily Laphriinae is a diverse taxon in the Afrotrop- tropics, comprising nine genera (13 genera globally), four of ics, comprising 33 genera (112 genera globally), 19 of which which are endemic and 132 species. This subfamily is global are endemic and 201 species. This subfamily is global in distri- in distribution, except Antarctica. Three of the four endem- bution, except Antarctica. Most genera (26) include less than ic genera have between 1–3 species each, while Afroestricus 10 species and the majority of species are placed in four gen- Scarbrough has 21. Scarbrough (2010) published an identifi- era, each with 22–29 species. Laphriinae was newly deline- cation key to Afrotropical genera and subgenera that has been ated by Dikow (2009a), who combined the Laphriinae and adjusted and incorporated into the below key. Laphystiinae sensu Geller-Grimm (2004) and Papavero (1973) as Laphystiinae was shown to be paraphyletic. The subfamily Stenopogoninae is a highly diverse taxon in the Afrotropics, comprising 22 genera (currently 28 genera The subfamily Leptogastrinae is a diverse taxon in the Afro- globally), 17 of which are endemic and 269 species. This tropics, comprising seven genera (18 genera globally), one of subfamily is global in distribution, except Antarctica. Sixteen which is endemic and 150 species. This subfamily is almost genera, 15 of which are endemic, have less than ten species global in distribution, being absent only from Chile, New Zea- and Gonioscelis Schiner with 40 species is the most diverse land and Antarctica. With the exception of Lasiocnemus Loew, Afrotropical endemic. The global genus Microstylum Macquart

Figs 48.42–45. Photographs of living Asilidae (Leptogastrinae): (42) Euscelidia sp. ♀ with araneaen prey (South Africa); (43) aganniphe Janssens ♂ with dipteran prey (South Africa); (44) Lobus sp. ♀ (Tanzania); (45) Mesoleptogaster sp. ♀ (Namibia). Fig. 42 (photograph © J.G.H. Londt), Fig. 43 (photograph © T. Dikow), Figs 44, 45 (photographs © S.A. Marshall).

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1108 SURICATA 5 (2017) with 79 Afrotropical species is one of the most diverse taxa and 27 species. Tillobromatinae only occurs in the Afrotropics, and occurs throughout the region. Stenopogoninae was newly where it has its largest species diversity and in the Neotrop- delineated by Dikow (2009a), as the monophyly of the previ- ics. Tillobromatinae was elevated to subfamily status by Dikow ous delineation sensu Artigas & Papavero (1988) and Geller- (2009a) and includes genera previously assigned to Stenopo- Grimm (2004) could not be established and genera are as- goninae sensu Geller-Grimm (2004) and Papavero (1973). signed to Bathypogoninae, Brachyrhopalinae (2 Afrotropical genera), Phellinae, Tillobromatinae (2) and Willistonininae (3). The subfamily Trigonomiminae is a medium-sized taxon, comprising two Afrotropical genera (12 genera globally), one The subfamily Stichopogoninae is a small taxon in the Afro- of which is endemic and 61 species. This subfamily has a glob- tropics, comprising six genera (14 genera globally), one of al distribution, with the exception of the Oceanian Region which is endemic and 26 species. This subfamily is global in and Antarctica. Oldroyd (1974) included Oligopogon Loew distribution, except Antarctica. Oldroyd (1963) included Psili­ in an identification key to the taxon and in 1980 catalogued nus Wulp and Sporadothrix Hermann in an identification key Oligopogon and Oxynoton Janssens in the tribe Xenomyzini to the taxon. Psilinus is a synonym of Rhabdogaster, placed in (within Stenopogoninae at that time). Oligopogon is placed as Brachyrhopalinae and Sporadothrix is placed in Willistonini- incertae sedis here, following Dikow (2009a) and Oxynoton is nae. Bosák & Hradský (2011: 739) recorded two genera for placed in the subfamily Brachyrhopalinae. the first time for the region. The subfamily Willistonininae is a small taxon, comprising The subfamily Tillobromatinae is a small taxon in the Afro- six Afrotropical genera (eight genera globally), five of which tropics, comprising two endemic genera (three genera globally) are endemic and 34 species. The subfamily is confined to

Figs 48.46–48. Photographs of living Asilidae (Ommatiinae): (46) Cophinopoda pulchripes (Bigot) ♀ (Réunion Is.); (47) sp. ♂ (Tanzania); (48) same ♀ (United Arab Emirates). Fig. 46 (photograph © D. Martiré), Fig. 47 (photograph © S.A. Marshall), Fig. 48 (photograph © H. Roberts).

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Figs 48.49–54. Photographs of living Asilidae (Stenopogoninae): (49) Daspletis vespertilio (Engel) ♂ (South Africa); (50) D. setitho­ racicus (Ricardo) ♀ (South Africa); (51) D. vulpes Loew ♂ (South Africa); (52) Gonioscelis sp. ♂ (Namibia); (53) Microstylum sp. ♀ (South Africa); (54) same ♂ with coreid prey (South Africa). Figs 49, 54 (photographs © J.G.H. Londt), Figs 50, 52 (photo- graphs © T. Dikow), Figs 51, 53 (photographs © S.A. Marshall).

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1110 SURICATA 5 (2017)

Figs 48.55–60. Photographs of living Asilidae (Stenopogoninae and Stichopogoninae): (55) Microstylum sp. ♀ (left) and ♂ (right) in copula, ♀ with hymenopteran prey (Namibia); (56) Scylaticus sp. ♀ (South Africa); (57) Rhadinus sp. ♀ (United Arab Emir- ates); (58) sp. ♀ with coleopteran prey (South Africa); (59) Stichopogon sp. ♂ (above) and ♀ (below) in copula (South Africa); (60) Stichopogon sp. ♀ with araneaen prey (Zambia). Figs 55, 56, 58, 59 (photographs © S.A. Marshall), Fig. 57 (photograph © H. Roberts), Fig. 60 (photograph © R. Felix).

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Figs 48.61–65. Photographs of living Asilidae (Tillobromatinae and Trigonomiminae): (61) Hypenetes stigmatias Loew ♂ (South Africa); (62) Damalis sp. ♀ (South Africa); (63) D. speciosa Loew ♀ (above, perching) and ♂ (below, upside-down) in copula (South Africa; see Londt 1991 for copulation sequence); (64) Rhipidocephala sp. ♂ (South Africa); (65) Rhipidocephala sp. ♀ (South Africa). Fig. 61 (photograph © J.G.H. Londt), Figs 62, 64 (photographs © S.A. Marshall), Figs 63, 65 (photographs © T. Dikow).

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1112 SURICATA 5 (2017) the Afrotropics (where it has its largest species diversity), the Identification south-western Palaearctic and western Nearctic Regions. Wil- listonininae was elevated to subfamily status by Dikow (2009a) Asilidae is the largest family of Afrotropical Diptera, currently and includes genera previously assigned to Stenopogoninae with 1,684 described species, representing about 8% of the sensu Artigas & Papavero (1988) and Geller-Grimm (2004). entire Diptera fauna (Kirk-Spriggs & Stuckenberg 2009: 175). The oldest assassin-fly descriptions from the Afrotropical Re- Taxa such as Asilinae, Dasypogoninae, Laphriinae, Lepto- gion were published by J.C. Fabricius and A.A.H. Lichtenstein gastrinae, Ommatiinae, Stichopogoninae and Trigonomiminae from West Africa and South Africa between 1775 and 1796, are no doubt monophyletic in the delineation used in the syn- i.e., Promachus fasciatus (F., 1775) (Sierra Leone, Asilinae), opsis below. Potential changes to taxon assignment are most Teratopomyia cyanea F., 1781 (South Africa, Brachyrhopal­ probable for Brachyrhopalinae, Stenopogoninae and Tillobro- inae), morio F., 1794 (“Guinea”, unplaced species), A. matinae, as the monophyly needs to be tested by including Lichtenstein, 1796 (South Africa, unplaced species), additional genera from the world fauna. A. melanotarsus Lichtenstein, 1796 (South Africa, unplaced species) and A. nigellus Lichtenstein, 1796 (South Africa, un- The morphological phylogenetic analysis of Dikow (2009a) placed species). The earliest comprehensive treatment of parts was unable to place the Afrotropical genus Oligopogon and of the Afrotropical fauna was published by Loew (1860) focus- Nearctic genus Coleomyia Wilcox & Martin in any of the 14 ing on the Southern African fauna. subfamily taxa and these are, therefore, placed as incertae se­ dis within Asilidae. Akatiomyia Londt is added here, as it is Considering the global fauna, assassin flies are known to morphologically similar to Oligopogon and new phylogenetic occur on all continents, except Antarctica and are diverse research is required to place both genera. on many islands/island groups, but are absent from

Figs 48.66–68. Photographs of living Asilidae (Willistonininae): (66) Acnephalomyia andrenoides (Wiedemann) ♀ (South Africa); (67) Sisyrnodytes sp. ♂ (United Arab Emirates); (68) Trichoura tankwa Londt ♀ (South Africa). Fig. 66 (photograph © S.A. Mar- shall), Fig. 67 (photograph © H. Roberts), Fig. 68 (photograph © T. Dikow).

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1113 the Hawaiian Is. The greatest species diversity is found in is Gonioscelis with 40 species followed by Rhabdogaster (39), semi-arid to arid environments and tropical rainforests. Londt Dasophrys Loew (32), Rhipidocephala Hermann (26), Proag­ (1998a) recorded 61 species from the semi-arid Karoo near onistes (22) and Hypenetes Loew and Afroestricus, each with Willowmore (Eastern Cape Province, South Africa) and Fisher 21 species. By far the most speciose genus in the Afrotropics (1985, 2009: 616) recorded 97 species from the Tambopata is Neolophonotus with 254 Afrotropical species, which is also Reserve in Amazonian Peru and 108 species from La Selva represented by two species in the Palaearctic Region and Ol- Biological Station in a Central American lowland rainforest droyd (1975: 147) disputes the presence of Neolophonotus in in Costa Rica. the Oriental Region. This genus is the most commonly encoun- tered assassin-fly taxon particularly in Southern Africa where its In an Afrotropical context, robber flies occur in almost every centre of diversity lies. biome, but appear to have their greatest diversity in arid hab- itats. Assassin flies have not been recorded from the South There are 22 described species that have not, for a variety Atlantic Ocean islands of Ascension, St. Helena, Tristan da of reasons, been assigned to any Afrotropical genus (Oldroyd Cunha and Gough, as well as the Cape Verde Is. Taxonomical- 1980: 373 lists 14 unplaced species). In all instances, the type ly, Afrotropical species are best known from arid to semi-arid specimens require re-examination and at least some of these Southern Africa, as well as the savanna belt from western Afri- appear to have been lost. A list of these species is provided ca through eastern Africa. The fauna of the tropical rainforest here for reference, along with the country/region from where in western and Central Africa and the arid environments of the they were originally described: Asilus bombylius Lichtenstein, southern Arabian Peninsula is much less known, due largely to 1796 (South Africa), A. forficula Macquart, 1846 (South Af- the paucity of material. Through the comprehensive revision- rica), A. gabonicus Macquart, 1855 (Gabon), A. incisuralis ary work of the senior author over the past 40 years the fauna Macquart, 1838 (South Africa), A. melanotarsus Lichtenstein, has been reviewed extensively and this is particularly true for 1796 (South Africa), A. morio F., 1794 (“Guinea”), A. natali­ Southern Africa. cus Macquart, 1855 (South Africa), A. nigellus Lichtenstein, 1796 (South Africa), A. schedius Walker, 1849 (South Africa), The Afrotropical Asilidae fauna is one of the most diverse in Dasypogon­ aequalis Walker, 1857 (South Africa), D. atripennis the world. Of the 557 genera globally, 148 genera (26.6%) oc- Macquart, 1834 (Senegal), D. caffer Wiedemann, 1828 (South cur within the region and 1,684 species (ca 22% of the world Africa), D. carvilius Walker, 1849 (unknown), D. fossius Walk- fauna) are known to date, with many additional species await- er, 1849 (South Africa), D. lenticepsa Thomson, 1869 (South ing description. Africa), D. reinhardi Wiedemann, 1824 (South Africa), D. trag­ Although the Afrotropical fauna has been catalogued (Ol- icus Wiedemann, 1828 (South Africa), stigmatizans F. , droyd 1980), no single comprehensive manuscript reviewing 1805 (Mauritius), Lophonotus albiciliatus Loew, 1854 (Egypt; the assassin-fly fauna has been published. Oldroyd’s (1970, Sudan), L. dubius Bezzi, 1892 (Somalia), L. leucotaenia Bez- 1974) reviews of regional faunas (Congo basin and Southern zi, 1906 (Eritrea) and L. macropterus Loew, 1854 (Nubien = Africa), complete with identification keys to genera, are among Sudan). the more useful general references for the region, in light of this contribution these are now outdated. Londt published a Londt (2002b) published the most recent key to the then rec- series of papers in the Annals of the Natal Museum entitled ognised subfamilies of Afrotropical Asilidae (i.e., Apocleinae, “Afrotropical Asilidae 1–33” between 1977 and 2000. In more Asilinae, Dasypogoninae, Laphriinae, Laphystiinae, Leptogas- than 55 additional manuscripts, the senior author reviewed trinae, Ommatiinae, Stenopogoninae, Stichopogoninae and many other genera and through contributions of other authors Trigonomiminae), which is superseded by the below identifi- many taxa were studied, however, a few of the larger global cation key. Although efforts have been made to avoid the use genera still require attention (e.g., Leptogaster Meigen, Micro­ of terminalia characters, this was not always possible and so, stylum and Promachus). The following eleven specialists have ideally, representatives of both sexes should be available when been the main contributors having each generated at least 50 the below key is used. The Asilinae genus Tolmerus still requires specific names (currently valid or otherwise) – J.G.H. Londt clarification in an Afrotropical context (see below) and is not (29% of fauna), H. Oldroyd (9%), H. Loew (6%), S.W. Bromley included in the below key. (6%), G. Ricardo (5%), J.P.M. Macquart (4%), F. Walker (3%), E. Janssens (3%), C.H. Martin (3%), E.O. Engel (3%) and A.G. The identification key provided below can also be accessed Scarbrough (3%). The majority of genera occurring in the Afro- electronically at https://asiloidflies.si.edu/content/online-iden- tropical Region have likewise been described by few dipter- tification-keys and http://keys.lucidcentral.org/keys/phoenix/ ists – J.G.H. Londt (44 genera, 30% of fauna, 42 endemic), H. Afrotropical_Asilidae_genera/. Loew (26, 18%, 11), F. Hermann (13, 9%, 6), H. Oldroyd (10, 7%, 10) and F.M. Hull (7, 5%, 5). The photographs of pinned specimens in Figs 85–236 rep- resenting 69 genera, plus additional views of the head in ante- Of the 148 genera in the Afrotropics, 91 (61.5%) are en- rior view, can be accessed electronically at Morphbank under demic to the region and 13 (8.8%) additional genera have their Collection ID 860613 (http://www.morphbank.net/myCollec- greatest species diversity and centre of distribution in the Afro- tion/?id=860613). tropics (four or less species known from outside the Afrotropi- cal Region). There are 31 (20.9%) endemic, monotypic genera Larger specimens of Asilidae should be direct-pinned and within the Afrotropical Region and 33 (22.4%) endemic gen- smaller species micro-pinned in the field for later staging (see era with 2–4 species each. The most diverse endemic genus Chapter 2).

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1114 SURICATA 5 (2017)

Key to genera of Afrotropical Asilidae

1. Abdominal tergite 2 not more than 4 × as long as wide (e.g., Fig. 17); abdominal sternite 1 confined beneath tergite 1; wing with alula (e.g., Fig. 84) and tarsi with pulvilli (e.g., Fig. 46) usually present, but occasionally one or other may be absent...... 2 – Abdominal tergite 2 at least 5 × or more as long as wide (e.g., Figs 42–45); abdominal sternite 1 extend- ing ca halfway back beneath tergite 2; wing without alula (e.g., Fig. 171) and tarsi without pulvilli (e.g., Fig. 42) (LEPTOGASTRINAE)...... 15 2. Fore tibia without spine-like tibial processes (macrosetae may be present); thorax with prosternum either fused to or separated from proepisternum by membranous area ...... 3 – Fore tibia with apical, unarticulated spur (i.e., a spine-like process of distal tibial margin, somewhat sigmoid in shape, e.g., Fig. 23); thorax with prosternum separated from proepisternum by membranous area (DASYPOGONINAE) ...... 21

3. Wing vein R2+3 ending in costal vein (C), cell r1 thus open at wing margin (as in Fig. 185) ...... 4

– Wing vein R2+3 joining vein R1 proximal to end of R1, cell r1 thus closed, either stalked or closed at wing margin (as in Figs 83, 84) ...... 10 4. Head with frons virtually same width at level of antennal insertion and vertex, or only slightly diverging 1 (e.g., Figs 74, 80); more or less oval or posterior margin slightly sinuate in ventral ∕4 (e.g., Figs 158, 224); thorax with prosternum either fused to or separated from proepisternum by membranous area ...... 5 – Head with frons narrowed at level of antennal insertion, then abruptly diverging dorsally (frons much wider at vertex than at antennal insertion) (e.g., Figs 58, 81); posterior margin of distinctly sinuate 1 in ventral ∕2 (e.g., Fig. 60); thorax with prosternum fused to proepisternum, forming a precoxal bridge (STICHOPOGONINAE) ...... 69 5. Antennal stylus with long setulae on all surfaces (either loosely arranged or tightly-packed) (e.g., Fig. 197); wing with alula reduced (e.g., Fig. 197)...... 6 – Antennal stylus without long setulae (e.g., Fig. 220, Rhipidocephala sometimes with ventral setulae); wing with alula usually well-developed (e.g., Figs 185, 186) ...... 7 6. Antennal stylus robust, broader than distal end of postpedicel; stylus setulae short and tightly-packed; thorax with prosternum separated from proepisternum by membranous area (confined to Western Cape Province, South Africa)...... Akatiomyia Londt – Antennal stylus narrower than distal end of postpedicel (Fig. 198); stylus setulae long and loosely ar- ranged (Fig. 198); thorax with prosternum fused to proepisternum, forming precoxal bridge (Fig. 198) (widespread throughout Afrotropical Region) ...... Oligopogon Loew

7. Head with tentorial pits well-developed, conspicuous anteroventrally (e.g., Fig. 71); wing cell m3 open (e.g., Figs 137, 215) (TRIGONOMIMINAE)...... 8

– Head with tentorial pits small, slit-like, inconspicuous ventrally (e.g., Fig. 78); wing cell m3 open or closed (e.g., Figs 163, 205)...... 9 8. Antennal postpedicel consisting of proximal bulb, with elongate, but undifferentiated distal part (resem- bling stylus) (Fig. 138); abdominal tergite 1 entirely sclerotised (Fig. 137); hind trochanter with posteriorly- directed, setose protuberance; fairly robust flies, often shiny yellow-brown with glistening (Figs 62, 63, 137, 138)...... Damalis F. – Antennal postpedicel elongate, with distinct apical stylus (Fig. 216); abdominal tergite 1 often unscle- rotised medially (Fig. 216); hind trochanter without protuberance; small, fragile, blackish flies (Figs 64, 65, 215, 216)...... Rhipidocephala Hermann 9. Female terminalia with divided tergite 10, bearing acanthophorite spines (may be difficult to discern in specimens with withdrawn terminalia) (e.g., Figs 189, 190, 306); thorax with anepisternum invar- iably without strong macroseta at superoposterior angle (e.g., Fig. 142); prosternum separated from proepisternum by membranous area ...... 23 – Female terminalia simple, tergite 10 never divided, without acanthophorite spines (e.g., Fig. 216); thorax with anepisternum usually with at least one macroseta at superoposterior angle, in front of wing insertion (e.g., Fig. 228) (if absent, then pulvilli also absent); prosternum fused to proepisternum, form- ing precoxal bridge (LAPHRIINAE [in part]) ...... 74

10. Wing vein R2+3 bent anteriorly at tip, joining vein R1 just before or at costal vein (C) (e.g., Figs 83, 233) ...... 11

– Wing vein R2+3 closing cell r1 by obvious, fairly straight stalk (e.g., Fig. 95) ...... 13

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1115

11. Costal wing vein (C) circumambient (as in Fig. 84); hind femur slender, without ventrodistal macrosetae inserted on tubercles (confined to eastern and southern Africa) ...... Perasis Hermann – Costal wing vein (C) terminating at or before vein CuA+CuP (cell cup not bordered by costal vein (C)) (e.g., Figs 83, 155, 233); hind femur stout, frequently with ventrodistal macrosetae inserted on tubercles (e.g., Figs 156, 234) (throughout sub-Saharan Africa and/or southern Arabia)...... 12 12. Antennal postpedicel almost 2 × as long as scape and pedicel combined (Fig. 155); larger species (> 13 mm); hind femur with ventrodistal macrosetae inserted on tubercles; wing with patches of yellow-brown infuscation (Fig. 155) (throughout sub-Saharan Africa); . . . .Hoplistomerus Macquart – Antennal postpedicel only slightly longer than scape and pedicel combined (Fig. 233); smaller species (< 13 mm); hind femur with or without ventrodistal macrosetae inserted on tubercles (Fig. 234); wing usually without infuscation (Fig. 233) (throughout sub-Saharan Africa and southern Arabia)...... Trichardis Hermann 13. Thorax with anepisternum invariably without strong macroseta at superoposterior angle (e.g., Fig. 142); prosternum either fused to or separated from proepisternum by membranous area; palpus 1-segmented ...... 14 – Thorax with anepisternum usually with at least one macroseta at superoposterior angle, in front of wing insertion (e.g., Fig. 228) (if absent, then pulvilli also absent); prosternum fused to proepisternum, form- ing precoxal bridge; palpus 1- or 2-segmented (LAPHRIINAE [in part])...... 78 14. Antennal stylus with long setulae (e.g., Fig. 146); thorax with postmetacoxal area sclerotised (i.e., post- metacoxal bridge complete) (OMMATIINAE)...... 102 – Antennal stylus without setulae (e.g., Fig. 204); thorax with postmetacoxal area unsclerotised and mem- branous at least medially (ASILINAE) ...... 110 15. Tarsal claws of unequal length, median claw shorter than lateral; wing veins M and R with trichoid spicules (reminiscent of setulae); thorax with long, posteriorly directed postsutural dorsocentral setae; male termi- nalia without surstylus (Fig. 281) (confined to south-western Madagascar)...... Schildia Aldrich – Tarsal claws of equal length, median and lateral claws equally long; wing veins M and R without trichoid spicules; thorax usually without posteriorly directed postsutural dorsocentral setae; male terminalia invariably with distinct surstylus (e.g., Figs 275, 278) (throughout Afrotropical Region)...... 16 16. Wing cell cua closed (as in Fig. 83)...... 17 – Wing cell cua open (as in Fig. 199)...... 18 17. Hind tibia enlarged, swollen, widest medially, with at least 2 long ventrally-angled setae on ventral sur- 3 face (Figs 167, 168); hind femur in distal ∕4 long setose on all surfaces (Figs 167, 168); wing membrane usually brown patterned (Fig. 167)...... Lasiocnemus Loew – Hind tibia cylindrical, usually widest at distal tip, without long ventrally-angled setae on ventral surface 3 (Fig. 104); hind femur in distal ∕4 not long setose (Fig. 104); wing membrane never brown patterned (although sometimes appearing darker due to dense microtrichia) (Fig. 103) ...... Ammophilomima Enderlein 18. Thorax with postpronotum medially with distinct, cylindrical, peg-like protuberance (Fig. 148) ...... Euscelidia Westwood – Thorax with postpronotum medially without protuberance (median postpronotum may be slightly ele- vated) (e.g., Figs 172, 174, 176, 182) ...... 19 19. Hind tibia with dorsal and ventral rows of short macrosetae (Figs 45, 181, 182); hind legs long, almost cylindrical throughout (Figs 45, 181, 182) ...... Mesoleptogaster Frey – Hind tibia without distinct rows of short macrosetae (single macrosetae may be present particularly towards distal tip) (e.g., Fig. 44); hind legs not markedly elongated, at least femur distinctly clubbed distally (e.g., Fig. 44) ...... 20 20. Antennal postpedicel short (as long as, or marginally longer than scape and pedicel combined); male terminalia with hypandrium and gonocoxite fused to form “lobus”; surstylus without movable second- ary ventral lobe, invariably bipartite distally; female terminalia often with long macrosetae distally on abdominal sternite 8...... Lobus Martin – Antennal postpedicel short or long (Figs 172, 174); male terminalia with hypandrium and gonocoxite invariably separated; surstylus invariably with movable secondary ventral lobe, with only single tip (sometimes more or less blunt with distinct dorso- and ventrodistal points); female terminalia without long macrosetae distally on abdominal sternite 8...... Leptogaster Meigen

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1116 SURICATA 5 (2017)

21. Fore tibial spur slender, sigmoid, not associated with proximal tarsal process, at most with group of stout, peg-like, black, tarsal macrosetae (Fig. 218); scutellum with 2 well-developed apical macrosetae (Fig. 217); male terminalia with epandrium and hypandrium unfused (suture distinctly visible)...... Loew – Fore tibial spur stout, associated with well-developed proximal tarsal process (e.g, Fig. 200); apical scutellar macrosetae highly reduced to absent (e.g., Fig. 200); male terminalia with epandrium and hypandrium fused, forming continuous ring (no suture apparent) (e.g., Fig. 262) ...... 22 22. Head with ocellar tubercle prominent (Fig. 200); median occipital sclerite with 2 vertical macrosetae; postpronotal lobe without macrosetae (Fig. 200); weak dorsocentral macrosetae may be present post- suturally...... Pegesimallus Loew – Head with ocellar tubercle not prominent; median occipital sclerite with 4–6 vertical macrosetae; post- pronotal lobe with 1–2 stout macrosetae; pre- and postsutural dorsocentral macrosetae present ...... Caroncoma Londt 23. Thorax with anatergite setose (e.g., Figs 108, 144, 186)...... 24 – Thorax with anatergite bare (e.g., Figs 118, 222) ...... 32 24. Antennal stylus composed of 2–3 distinctly discernible articles (1–2 segment-like articles plus apical seta-like sensory article) ...... 25 – Antennal stylus reduced, composed of greatly reduced segment-like articles (not always evident) and apical seta-like sensory article in cavity on distal postpedicel ...... 31 25. Head with occiput without macrosetae (i.e., with weak postocular and occipital setae only)...... 26 – Head with occiput with obvious macrosetae (including postocular and occipital setae) (e.g., Fig. 108) ...... 27 26. Antennal scape distinctly longer than pedicel; eye : face width ratio < 1.1 : 1; male terminalia with 1 hypandrium less than ∕2 as long as epandrial lobes...... Dioctobroma Hull – Antennal scape and pedicel of virtually same length; eye : face width ration > 1.3 : 1; male terminalia with hypandrium ca as long as epandrial lobes...... Dogonia Oldroyd 27. Thorax with proepisternum with a few strong macrosetae in addition to setae (Fig. 108); antepronotum and scutum with very strong macrosetae, giving a bristly appearance (Fig. 108). . .Anasillomos Londt – Thorax with proepisternum with setae only (some may be stronger than others, but never as strong as scutal macrosetae); antepronotum and scutum with setae or regular macrosetae...... 28 28. Abdominal tergite 1–4 with group of strong macrosetae dorsolaterally; antennal stylus composed of two articles (1 segment-like article, plus apical seta-like sensory article); male terminalia bulbous (Figs 295–297); female abdominal tergites 7 and 8 of virtually same length. . . Ontomyia Dikow & Londt – Abdominal tergite 1 (only) with group of strong macrosetae dorsolaterally; antennal stylus composed of three articles (2 segment-like articles, plus apical seta-like sensory article); male terminalia slender, never bulbous; female abdominal tergite 8 distinctly shorter than tergite 7 (Fishermyia female unknown) ...... 29 29. Head with facial swelling well-developed on entire face...... Oratostylum Ricardo – Head with facial swelling weak, only ventral margin moderately developed ...... 30

1 30. Scutellum with 2 apical scutellar macrosetae; head with mystax occupying ventral ∕3 of face only; an- tennal postpedicel elongate, spindle-shaped, > 1.5 × as long as scape and pedicel combined (confined to Madagascar) ...... Fishermyia Londt – Scutellum with 4 or more apical scutellar macrosetae present; head with mystax occupying almost entire face, although sometimes weak in dorsal part; antennal postpedicel usually somewhat clavate, < 1.5 × as long as scape and pedicel combined (confined to southern Africa)...... Remotomyia Londt

3 31. Head with facial swelling occupying ca ∕4 of face (Fig. 144); mystax entirely covering face (Fig. 144); thoracic pre- and postsutural dorsocentral setae well-developed; postmetacoxal membrane clothed in

long setae (Fig. 144); wing vein M1 not strongly arched anteriorly (Fig. 143) ...... Daspletis Loew 1 – Head with facial swelling occupying at most ventral ∕2 of face (Figs 186, 188, 190); mystax primarily 1 confined to ventral ∕2 of face (Figs 186, 188, 190); only postsutural dorsocentral setae well-developed;

postmetacoxal membrane usually bare (Figs 186, 188, 190); wing vein M1 usually strongly arched ante- riorly (Figs 185, 187, 189) ...... Microstylum Macquart

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32. Costal wing vein (C) not circumambient, cell cup and alula without bordering vein C (i.e., C terminates at or before point where vein CuP joins wing margin) (e.g., Figs 91, 93, 105, 117, 211, 221, 223). . . 33 – Costal wing vein (C) circumambient (around entire wing margin, even when weakly-developed as in Trichoura) (e.g., Fig. 157)...... 43 33. Tarsal claws with pulvilli minute or absent (e.g., Fig. 224) (WILLISTONININAE [in part]) ...... 34 – Tarsal claws with pulvilli well-developed (e.g., Fig. 212) ...... 38 34. Abdomen broad, obviously dorsoventrally flattened (width : length ratio of abdominal tergite 2 > 2) (e.g., Figs 85, 222)...... 35 – Abdomen cylindrical, not obviously dorsoventrally flattened (width : length ratio of abdominal tergite 2 < 1.5) (e.g., Figs 102, 224) ...... 37

35. Costal wing vein (C) terminating before reaching vein CuA+CuP (e.g., Fig. 221); cell r5 closed and stalked (stalk frequently not reaching wing margin) (e.g., Fig. 221)...... Sisyrnodytes Loew – Costal wing vein (C) terminating at point where vein CuA+CuP reaches wing margin (e.g., Fig. 85); cell

r5 open (e.g., Fig. 85)...... 36

36. Wing cell m3 open (Fig. 85); vein R4 with short stump vein (Fig. 85); tarsal claws with pulvilli poorly developed, but clearly discernible; male terminalia with hypandrium somewhat flat or only gently con- cave...... Acnephalomyia Londt

– Wing cell m3 closed and stalked; vein R4 without stump vein (Fig. 117); tarsal claws with pulvilli minute, difficult to discern; male terminalia with hypandrium distinctly cup-shaped ...... Astiptomyia Londt 37. Small species (wing length: < 3.5 mm) (Fig. 101); tarsal claws without empodium (Fig. 102); wing

without stump vein on vein R4; thorax with scutal macrosetae well-developed, many times longer than accompanying setae...... Ammodaimon Londt – Larger species (wing length:> 6.0 mm) (Fig. 223); tarsal claws with well-developed empodium (Fig.

224); wing with short stump vein on vein R4 (Fig. 223); thorax with scutal macrosetae moderately de- veloped, not considerably longer than accompanying setae (Fig. 224) . . . . Sporadothrix Hermann 38. Thoracic postmetacoxal area sclerotised (i.e., postmetacoxal bridge complete) (Fig. 212)...... Rhabdogaster Loew – Thoracic postmetacoxal area entirely membranous ...... 39 39. Thorax with scutum greatly elevated anterodorsally, hump-like ...... Oxynoton Janssens – Thorax with scutum of more usual shape, not hump-like...... 40 40. Hind coxa anteriorly with elongate, distally rounded, peg-like process ...... Ischiolobos Londt – Hind coxa anteriorly without peg-like process...... 41 41. Head with strong mystacal macrosetae extending over entire face (Fig. 106); thorax with 1–4 long post- alar macrosetae (Fig. 106) ...... Amphisbetetus Hermann

1 – Head with strong mystacal macrosetae either confined to lower ∕2 of face or absent (weak mystacal setae present) (e.g., Figs 92, 94); thoracic postalar macrosetae usually absent (1 or 2 macrosetae in some species) (e.g., Figs 92, 94)...... 42 42. Head with mystacal macrosetae more extensive, if restricted to single row on lower facial margin, then weaker mystacal setae also present on face (e.g., Figs 92, 94); abdominal tergites usually bare (Figs 91–94) (primarily in Southern Africa)...... Afroholopogon Londt – Head with mystacal macrosetae restricted to single row on lower facial margin; abdominal tergites pri- marily pubescent (small median bare maculae present) (Palaearctic & southern Arabian Peninsula) ...... Wadipogon Bosák & Hradsky 43. Palpus 1-segmented (usually fairly robust and curved, such that their distal ends converge) . . . . 44 – Palpus 2-segmented (weak distal segment may be tucked away in oral cavity)...... 46 44. Proboscis with spine-like processes distally; antennal stylus not clearly differentiated, apical seta-like sensory article situated sub-apically in cavity on postpedicel...... Hynirhynchus Lindner – Proboscis of more usual form, without spine-like processes distally (e.g., Figs 152, 208); antennal stylus clearly differentiated, apical seta-like sensory article situated apically in cavity on stylus...... 45

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1118 SURICATA 5 (2017)

1 45. Proboscis shorter than antenna (Fig. 152); mystax occupying ventral ∕3 of face (Fig. 152); male mid tarsomeres 4–5 with highly modified, rather spade-shaped setae (usually reddish in colour) (Fig. 152) ...... Habropogon Loew 1 – Proboscis longer than antenna (Fig. 208); mystax occupying ventral ∕2 of face (Fig. 208); male mid tar- someres 4–5 with regular setae only (Fig. 208) ...... Pycnomerinx Hull 46. Head almost circular in anterior view (i.e., face narrow) (e.g., Fig. 69) ...... 47 – Head distinctly wider than high in anterior view (e.g., Fig. 70)...... 51 47. Thorax with katatergite bare...... 48 – Thorax with katatergite setose (e.g., Fig. 98) ...... 49 48. Fore femur with large, proximoventral spinose process (i.e., strongly raptorial) (Figs 52, 150) ...... Gonioscelis Schiner – Fore femur of more usual form, without large spinose process...... Loew 49. Thoracic dorsocentral macrosetae developed pre- and postsuturally (Fig. 154) ...... Haroldia Londt – Thoracic dorsocentral macrosetae developed only postsuturally (e.g., Fig. 98) ...... 50 50. Antennal stylus composed of two articles (1 segment-like article, plus apical seta-like sensory article) (Fig. 98); male terminalia with hypandrium usually bifurcate distally . . . . . Afroscleropogon Londt – Antennal stylus composed of three articles (2 segment-like articles plus apical seta-like sensory article); male terminalia with hypandrium usually simple ...... Rhacholaemus Hermann 51. Proboscis strongly downwardly-curved, resembling a parrot’s beak (Fig. 110) ...... Ancylorhynchus Berthold – Proboscis of more usual form, not strongly downwardly-curved (e.g., Fig. 134)...... 52 52. Head with facial swelling strongly projecting ventrally only (not dorsally), giving pointed, nose-like ap- pearance; mystacal macrosetae largely confined to small area at apex of facial swelling...... Lycostommyia Oldroyd – Head with facial swelling not projecting ventrally, of more usual form; mystacal macrosetae placed more widely on facial swelling (Figs 134, 136)...... 53 53. Head with mystax occupying entire face, even if weakly-developed dorsally (e.g., Fig. 134) ...... 54 – Head with mystax not occupying entire face, with distinct gap between dorsal mystacal setae and an- tennal sockets (e.g., Fig. 220) ...... 59 1 54. Antennal postpedicel widening toward middle (in lateral view), apical ∕2 appearing strongly incised ventrally (P. astroptica Londt, 1994) ...... Pedomyia Londt [in part] – Antennal postpedicel either distinctly laterally compressed or cylindrical...... 55 55. Antennal postpedicel distinctly laterally compressed, strap-like; stylus laterally compressed...... Hermannomyia Oldroyd – Antennal postpedicel more or less cylindrical (may appear slightly laterally compressed) (Figs 133, 134); stylus more or less cylindrical ...... 56 56. Large, -mimicking flies (wing length: > 15 mm) (Figs 119, 120); palpus large, well-developed (Fig. 120); thoracic anepimeral macroseta absent (Fig. 120)...... Londt – Small flies (wing length: < 10 mm) (e.g., Figs 129, 130, 133–136); palpus moderately developed (e.g., Figs 130, 134, 136); thoracic anepimeral macroseta usually present (e.g., Figs 130, 136). . . . . 57 57. Male terminalia club-like (Fig. 134); epandrium greatly developed, hemispherical; hypandrium greatly reduced...... Corymyia Londt – Male terminalia of more usual form (e.g., Fig. 130); epandrium not greatly developed; hypandrium not greatly reduced...... 58 58. Head with mystax well-developed, occupying entire face (Fig. 130); scutellum with numerous apical scutellar setae, usually some discal scutellar setae (central area usually bare) (Fig. 129); male terminalia with gonocoxite with two sub-equal, pointed, distal processes, lateral one with at most a small tumid dorsodistal projection ...... Connomyia Londt – Head with mystax moderately developed, occupying entire face, but usually weak dorsally; scutellum usually with few apical scutellar setae, rarely few discal scutellar setae present; male terminalia with gonocoxite with lateral process having distal or dorsodistal flange-like process. . . .Danomyia Londt

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1119

59. Thorax with anepimeral macroseta; hind tarsus with empodium laterally compressed and blade-like...... Empodiodes Oldroyd – Thorax without anepimeral macroseta; hind tarsus with empodium seta-like, not laterally compressed and blade-like...... 60 3 60. Head with facial swelling strongly developed in ventral ∕4, dorsal part distinctly defined (e.g., Fig. 158) ...... 61 – Head with facial swelling at most moderately developed, dorsal part not distinctly defined (e.g., Fig. 220)...... 62 61. Body not entirely metallic blue-black (Fig. 158); antennal postpedicel strongly club-shaped, subequal to length of scape and pedicel combined (Fig. 158); wing largely transparent with dark maculae (especially

“pterostigma-like” marking at base of cell r1) (Fig. 157)...... Hypenetes Loew – Body entirely metallic blue-black; antennal postpedicel elongate, cylindrical, ca 2 × as long as scape and pedicel combined; wing fairly uniformly black ...... Teratopomyia Oldroyd 1 62. Head with mystax occupying at most ventral ∕3 of face (e.g., Figs 180, 236) ...... 63 1 – Head with mystax occupying at least ventral ∕2 of face (e.g., Fig. 220) ...... 67

63. Wing cells cua and m3 closed and stalked (Fig. 235); male terminalia with hypandrium reduced, largely fused with gonocoxites (WILLISTONININAE [in part])...... Trichoura Londt

– Wing cells cua and m3 open at wing margin (even if only narrowly); male terminalia with hypandrium moderately well-developed, not fused with gonocoxites ...... 64 64. Male terminalia with epandrial lobes separated, only joined proximally ...... 65 1 – Male terminalia with epandrial lobes fused medially for at least proximal ∕2 of length...... 66 65. Larger flies (wing length: > 5 mm) ...... Antiscylaticus Londt – Smaller flies (wing length: < 5 mm) ...... Microphontes Londt 66. Scutellum with discal setae (ca 4); male terminalia with epandrial lobes fused medially for virtually en- tire length (only slight distal indentation) ...... Irwinomyia Londt 1 – Scutellum without discal setae (Fig. 180); male terminalia with epandrial lobes fused medially for ca ∕2 length ...... Macroetra Londt 1 67. Antennal postpedicel widening toward middle (in lateral view), apical ∕2 appearing strongly incised ventrally...... Pedomyia Londt [in part] – Antennal postpedicel spindle-shaped (e.g., Fig. 219)...... 68 1 68. Male terminalia with epandrial lobes short, fused medially for ca ∕3 length; male hypandrium elongate, ventrally directed with upturned distal region...... Agrostomyia Londt – Male terminalia with epandrial lobes long, entirely separated medially or very narrowly joined proxi- mally; male hypandrium more or less straight and distally directed...... Scylaticus Loew 69. Tarsi without pulvilli ...... 70 – Tarsi with pulvilli (even if only weakly developed) ...... 71

70. Wing cell m3 open (Fig. 213); head with mystacal setae dorsoventrally flattened (Fig. 214)...... Rhadinus Loew

– Wing cell m3 closed; head with mystacal setae circular in cross-section . . .Turkmenomyia Paramonov 1 71. Tarsi with pulvilli weakly developed, about ∕2 length of tarsal claw (confined to Kenya) ...... Nanoculcita Londt & Copeland – Tarsi with pulvilli well-developed, almost reaching distal end of tarsal claw...... 72

72. Head with 2 well-developed ocellar macrosetae (Fig. 81); wing cell m3 with long stalk at base (Fig. 225); usually small, or even tiny flies (body length: < 10 mm) (Figs 58–60, 225, 226); habitat sandy banks of streams, rivers and lakes...... Stichopogon Loew

– Head with ocellar macrosetae weak or absent (e.g., Fig. 126); wing cell m3 with short stalk or stalk absent (e.g., Fig. 125); habitat not as above...... 73 73. Thorax with scutum entirely grey pubescent (Fig. 125); larger flies (body length: > 10 mm) (littoral zones on East African coast and Indian Ocean islands) ...... Clinopogon Bezzi – Thorax with scutum bare; smaller flies (body length: < 10 mm) (confined to southern Arabian Peninsula) ...... Dichropogon Bezzi

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1120 SURICATA 5 (2017)

1 74. Tarsi with pulvilli poorly developed (ca ∕2 length of tarsal claw) (Fig. 206). . . . . Prytanomyia Özdikmen – Tarsi with pulvilli well-developed (as long as or slightly shorter than tarsal claw) (e.g., Fig. 164) . . . . .75 75. Thorax with anepisternum with strong macroseta on superoposterior angle; small flies (body length: < 8 mm); head with face narrower than width of one eye in anterior view; antennal scape 2 × as long as pedicel ...... Gerrolasius Hermann – Thorax with anepisternum without obvious strong macroseta on superoposterior angle; larger flies (body length: > 8 mm); head with face as wide or wider than width of one eye; antennal scape < 2 × as long as pedicel...... 76 76. Head with mystax composed almost entirely of fine setae, uniformly covering face; fine setae of thorax

and abdomen longish and soft; no conspicuous macrosetae on body; wing cell r5 usually closed; bee- like in appearance ...... Pilophoneus Londt – Head with mystax composed of strong macrosetae, largely restricted to ventral facial margin (e.g., Fig. 164); fine setae of thorax and abdomen minute; thoracic and abdominal macrosetae conspicuous and

moderately well-developed (e.g., Fig. 164); wing cell r5 open (e.g., Fig. 163); not bee-like in appear- ance ...... 77 77. Scutellum with apical macrosetae moderately developed; antennal postpedicel virtually as long as scape and pedicel combined; thorax and abdomen extensively bare...... Ericomyia Londt – Scutellum without apical macrosetae (Fig. 163); antennal postpedicel considerably longer than scape and pedicel combined (Fig. 164); thorax and abdomen entirely setose (Figs 163, 164). . . . Laphyctis Loew 78. Thorax with postmetacoxal area sclerotised (i.e., postmetacoxal bridge complete, no suture evident) ...... 79 – Thorax with postmetacoxal area membranous (some Proagonistes with almost complete bridge, but dorsoventral suture evident)...... 84 79. Larger flies (body length: > 10 mm) (confined to Madagascar) ...... 80 – Small to tiny flies (body length: < 10 mm) (confined to sub-Saharan Africa)...... 81 80. Male terminalia with cercus short, barely extending beyond tip of epandrium. . . Katharma Oldroyd – Male terminalia with cercus very long, extending well beyond tip of epandrium ...... Katharmacercus Tomasovic 1 81. Antennal postpedicel without stylus, with seta-like sensory article inserted just beyond ∕2 length; thorax with scutum without distinct macrosetae anteriorly; anatergite bare; scutellum with tubercular projec- tion apically ...... Afromosia Londt – Antennal postpedicel with distinct stylus, seta-like sensory article inserted sub-apically and laterally on postpedicel; thorax with scutum anteriorly with pair of small macrosetae; anatergite setose; scutellum simple, smoothly rounded...... 82 82. Antennal stylus as long as or longer than scape; head with postocular macrosetae weakly developed ...... Dichaetothyrea de Meijere – Antennal stylus shorter than scape; head with postocular macrosetae well-developed ...... 83 83. Head with face wide (eye : face width ratio < 1.5); mystax of males with dorsoventrally flattened, shiny scale-like macrosetae (Fig. 28); thorax with scutum shiny, at most slightly puncticulate (Fig. 28)...... Goneccalypsis Hermann – Head with face narrow (eye : face width ratio > 1.5); mystax of both sexes without laterally situated dorsoventrally flattened scale-like setae; thorax with scutum punctate. . . . . Loewinella Hermann

84. Wing vein M3 with apical portion perfectly aligned with proximal portion of vein M2 (forming cross) (confined to Madagascar)...... Orthogonis Hermann – Wing veins not so aligned (e.g., Figs 111, 123, 159, 227) (except in Anypodetus nigrifacies Ricardo, 1925, in which alignment is almost perfect) (widespread Afrotropical Region) ...... 85 85. Proboscis long, narrow and laterally compressed (knife-like) (as in Fig. 124) ...... 86 – Proboscis short to moderate in length, often stout, somewhat triangular in cross-section (e.g., Fig. 228) ...... 88 86. Hind femur considerably expanded mediodistally and with ventrodistal macrosetae inserted on tuber- cles...... Storthyngomerus Hermann – Hind femur not obviously expanded mediodistally, with only regular ventrodistal macrosetae (not in- serted on tubercles) (e.g., Fig. 124)...... 87

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1121

87. Antennal postpedicel at most 2 × as long as scape and pedicel combined (Fig. 124); female ovipositor short and not markedly tubular; smaller (body length: < 20 mm), not obviously robust, setaceous, or bee-like flies (Figs 27, 123, 124)...... Choerades Walker – Antennal postpedicel > 2 × as long as scape and pedicel combined; female ovipositor somewhat elon- gate, tubular; large (body length: > 20 mm), robust, setaceous, bee-like flies...... Dasyllina Bromley 88. Palpus laterally compressed and leaf-like (e.g., Fig. 160); female ovipositor projecting distally as slender tube (e.g., Fig. 159)...... 89 – Palpus cylindrical in cross-section (e.g., Fig. 228); female ovipositor short, not markedly tubular . . . . 92

89. Wing cell r5 closed (Fig. 159); broad flies, bee-like in appearance (mimicking carpenter ) (Figs 159, 160)...... Hyperechia Schiner

– Wing cell r5 open; not broad and bee-like in appearance...... 90 90. Head with facial swelling only weakly developed; antennal postpedicel virtually as long as scape and pedicel combined...... Rondani – Head with facial swelling well to strongly developed; antennal postpedicel ca 1.5 × as long as scape and pedicel combined ...... 91 91. Larger flies (body length: > 20 mm) mimicking pompilid wasps (Figs 36, 37); thorax with scutum black- ish (sometimes with red-brown lateral parts) (Fig. 37)...... Proagonistes Loew – Smaller flies (body length: < 20 mm); thorax with scutum brown-yellow to reddish, clothed in golden setae...... Systropalpus Hull

92. Wing vein M2 not reaching wing margin...... Ctenota Loew

– Wing vein M2 reaching wing margin (e.g., Figs 161, 169, 227)...... 93 93. Head with postgena well-developed, with ventral flange-like projection (in lateral view) (e.g., Fig. 162) ...... 94 – Head with postgena simple, not ventrally extended and flange-like (e.g., Figs 112, 166, 170, 192). . . . 95 94. Hind leg greatly elongate (hind femur ca 2 × as long as mid femur) (Fig. 162); abdomen somewhat 1 constricted in anterior ∕2 (Fig. 161); larger flies (body length: ca 20–36 mm) (Figs 30, 161, 162)...... Lamyra Loew – Hind leg normally proportioned (hind femur not more than 1.5 × as long as mid femur) (Fig. 228); abdomen more or less parallel-sided, not noticeably constricted (Fig. 227); smaller flies (body length: ca 10–27 mm) (Figs 39, 227, 228)...... Stiphrolamyra Engel

95. Wing cell r5 closed (e.g., Fig. 169) ...... 96

– Wing cell r5 open (e.g., Figs 111, 165, 192)...... 98 96. Palpus bulbous (virtually spherical); mystax composed of fine setae only ...... Afromelittodes Oldroyd & Van Bruggen – Palpus not markedly bulbous (cylindrical) (e.g., Fig. 170); mystax composed of strong macrosetae (e.g., Fig. 170) ...... 97 97. Antennal postpedicel with numerous well-developed dorsal setulae (Fig. 169); scape usually ca 2 × as long as pedicel (Fig. 169); often rather bee-like in appearance (Figs 32, 169, 170) ...... Laxenecera Macquart – Antennal postpedicel without setulae; scape only slightly longer than pedicel; never bee-like in appear- ance ...... Nusa Walker 98. Tarsi without pulvilli (Fig. 112); thorax with anepisternum without macrosetae on superoposterior angle (Fig. 112) ...... Anypodetus Hermann – Tarsi with pulvilli (e.g., Figs 166, 192); thorax with anepisternum with strong macrosetae on superopos- terior angle (Figs 166, 192)...... 99 99. Antennal postpedicel bearing well-defined segment-like stylus, tipped with terminal pit-enclosed seta-like sensory article (Fig. 165)...... Laphystotes Oldroyd – Antennal postpedicel simple, tipped with terminal, obliquely positioned pit-enclosed seta-like sensory article (e.g., Fig. 191)...... 100 100. Generally pale yellow-brown to red-brown flies; legs entirely yellow-brown to red-brown...... Smeryngolaphria Hermann

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1122 SURICATA 5 (2017)

– Generally dark red-brown to black flies; legs extensively or entirely blackish (e.g., Figs 191, 192). . .101 101. Head of male with mystax at least laterally overlaid by shiny scale-like setae (Fig. 34); wing with proxi- 1 1 mal ∕2 transparent, distal ∕2 uniformly clothed in dense microtrichia...... Notiolaphria Londt – Head in both sexes with mystax not overlaid by shiny scale-like setae (Fig. 192); wing membrane more extensively clothed in black microtrichia (Fig. 192)...... Nannolaphria Londt 102. Head with mystax with some dorsal macrosetae arranged in two distinct vertical rows (e.g., Figs 47, 48, 90)...... 103 – Head with mystax simple, dorsal setae not arranged in distinct vertical rows (e.g., Fig. 46, 130, 184) ...... 108 103. Scutellum without apical macrosetae; antennal stylus with ventral setulae arranged in single row ...... Metommatius Hull – Scutellum with apical macrosetae (e.g., Figs 89, 146); antennal stylus with ventral setulae arranged in two distinct divergent rows (e.g., Figs 90, 146) ...... 104 1 104. Head with facial swelling moderately well-developed (at least in ventral ∕2) (e.g., Fig. 90)...... 105 – Head with facial swelling only weakly developed ventrally (e.g., Fig. 146) ...... 106 2 105. Head with facial swelling moderately well-developed, abruptly produced in ventral ∕3 (Fig. 90); pro- boscis more or less cylindrical medially (only slightly higher than wide), shorter than (in lateral view) (Fig. 90); antennal postpedicel small, virtually as wide as scape (Fig. 89); thorax with 2–3 stout, long presutural dorsocentral macrosetae (Fig. 89)...... Afroestricus Scarbrough 1 – Head with facial swelling less developed, with only ventral ∕2 developed; proboscis oval in diameter medially (higher than wide), as long as or slightly longer than compound eye (in lateral view); antennal postpedicel wider than scape; thorax with short presutural dorsocentral macrosetae ...... Longibeccus Scarbrough 106. Abdominal tergites 2–4 strongly constricted laterally (Fig. 145) ...... Emphysomera Schiner – Abdomen tergites 2–4 more or less parallel-sided (not noticeably constricted) (e.g., Figs 47, 48) . . . . . 107 107. Thorax with stout macroseta on anepimeron; male mystax usually narrow, individual macrosetae taper- ing evenly from proximal to distal; male abdominal sternites 3–4 without pattern of erect macrosetae and/or dense setae (Figs 47, 48)...... Ommatius Wiedemann – Thorax usually without macroseta on anepimeron, if present, then only setose, rarely macrosetose; male mystax usually with several thick macrosetae of uniform diameter medially, tapering only at or just before apex; male abdominal sternites 3–4 with pattern of erect macrosetae and/or dense setae ...... Pygommatius Scarbrough & Marascia 1 108. Head with face narrow at antennal insertion, < ∕5 as wide as head at greatest width; antennal postpedi- cel 2–6 × as long as scape and pedicel combined (Fig. 184); stylus short, much shorter than postpedicel (Fig. 184) ...... Michotamia Macquart 1 – Head with face wide, > ∕5 as width of head; antennal postpedicel short, virtually as long as scape or scape and pedicel combined (Fig. 46); stylus long, usually > 3 × as long as postpedicel (e.g., Fig. 46) ...... 109 109. Head with mystax composed of dense macrosetae medially (Figs 46, 132); numerous ocellar setae po- sitioned fan-like posteriorly; antennal stylus with setulae short, arranged in single row (Fig. 132); larger robust flies (body length: > 10 mm) (Figs 46, 131, 132) ...... Cophinopoda Hull – Head with mystax composed of sparse setae medially; only 2 ocellar setae posteriorly; antennal stylus with setulae long, arranged in two rows; smaller flies (body length: < 9 mm)...... Thallosia Oldroyd 110. Thorax with anatergite bare (although setae may be present on mediotergite (mesopostnotum)) (e.g., Figs 100, 114, 142, 178, 194, 196, 202, 204, 230, 232)...... 111 – Thorax with anatergite setose (e.g., Figs 96, 128)...... 136 111. Antennal stylus composed of two articles (one segment-like article plus apical seta-like sensory article); head with facial swelling weakly developed ventrally (e.g., Figs 100, 114, 202, 204); dorsocentral mac- rosetae only postsuturally (e.g., Figs 100, 114, 178, 202, 204)...... 112 – Antennal stylus composed of three articles (two unequal segment-like articles plus apical seta-like sen- sory article); head with facial swelling not distinctly defined or gently convex (Figs 17, 142, 194, 196, 230, 232); thorax with dorsocentral macrosetae usually inserted pre- or postsuturally (e.g., Figs 142, 194, 196, 230, 232) ...... 118

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1123

112. Wing with complete auxiliary vein on vein R4 (connecting veins R2+3 and R4) (as in Fig. 84; Figs 99, 201, 203)...... 113

– Wing either without auxiliary vein on vein R4 or with only short stump vein on vein R4 (e.g., Figs 113, 178)...... 116

113. Wing either with auxiliary vein long, or running parallel to vein R4+5 for considerable distance before reaching fork of veins R4 and R5 (e.g., Figs 84, 203); cell r4 short, diverging rapidly near wing margin (e.g., Fig. 203); posterior wing margin with microtrichia arranged in single row (aligned in same plane as wing membrane); scutellum with 8 or more apical macrosetae (e.g., Fig. 203); discal scutellar setae and macrosetae present (Fig. 204)...... 114

– Wing either with auxiliary vein shorter, or at most running parallel to vein R4+5 for short distance (e.g., Figs 99, 201); cell r4 long, diverging gradually towards wing margin (e.g., Figs 99, 201); posterior wing margin with microtrichia arranged in two rows (diverging from plane of wing membrane); scutellum with fewer than 8 apical macrosetae (e.g., Figs 99, 201); discal scutellar setae composed of setae only (no macrosetae) (e.g., Figs 99, 201) ...... 115 114. Head with vertex of regular width, with ocellar tubercle of regular shape and size (Figs 77, 203); ocellar tubercle usually not visible in lateral view (Fig. 204) ...... Promachus Loew – Head with vertex wide, entirely occupied by ocellar tubercle; ocellar tubercle distinctly visible in lateral view ...... Tuberconspicus Tomasovic

115. Wing with auxiliary vein very short (virtually as long as section of vein R4 between fork of vein R4+5 and point at which crossvein joins R4) (Fig. 99)...... Alcimus Loew – Wing with auxiliary vein longer (Fig. 201)...... Philodicus Loew

116. Wing without stump vein on vein R4 (Fig. 178); female ovipositor telescopic, comprised of abdominal segments 5 and following (confined to Madagascar & Mauritius) ...... Lycoprosopa Hull

– Wing with short stump vein (st vn) on vein R4 (e.g., Fig. 113); female ovipositor not obviously telescopic, comprising abdominal segments 8 and following (confined to West Africa, Arabian Peninsula & North Africa)...... 117

117. Scutellum with 2 apical macrosetae (Figs 113, 115); hind coxa with 2–3 lateral macrosetae; wing cell r5 open or closed (Figs 113, 115); female ovipositor tubular; cercus spinose with well-developed macro- setae...... Apoclea Macquart

– Scutellum with 4–6 apical macrosetae; hind coxa with single lateral macroseta; wing cell r5 open; fe- male ovipositor laterally compressed; cercus without macrosetae...... Erax Scopoli 118. Head with dorsal postocular setae short to moderately long, at most weakly proclinate (e.g., Fig. 232) ...... 119 – Head with dorsal postocular setae long, markedly proclinate (e.g., Figs 12, 142, 194, 196, 230). . . . . 123 119. Antennal stylus distinctly longer than postpedicel (e.g., Fig. 232) ...... 120 – Antennal stylus shorter or virtually same length as postpedicel...... 121 120. Thorax with postpronotal lobes with macrosetae in addition to fine setae; hind coxa with single lateral macroseta; posterior wing margin with microtrichia arranged in two rows (diverging from plane of wing membrane)...... Dysclytus Loew – Thorax with postpronotal lobes with fine setae only (Fig. 232); hind coxa with 4–5 lateral macrosetae; posterior wing margin with microtrichia arranged in single row (lying in same plane as wing mem- brane)...... Torasilus Londt 121. Thorax with dorsocentral macrosetae well-developed pre- and postsuturally; hind coxa with more than 2 lateral macrosetae...... Zelamyia Londt – Thorax with dorsocentral macrosetae well-developed only postsuturally; hind coxa with 2 lateral mac- rosetae...... 122 122. Female cercus with fine setae only (without dorsodistal projection) ...... Acasilus Londt – Female cercus with fine setae and dorsodistal spine-like projection...... Juxtasilus Londt 123. Posterior wing margin with microtrichia arranged in 2 rows (diverging from plane of wing membrane) ...... Robertomyia Londt – Posterior wing margin with microtrichia arranged in single row (lying in same plane as wing mem- brane)...... 124

124. Wing cell r5 closed well before wing margin ...... Megadrillus Bigot

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1124 SURICATA 5 (2017)

– Wing cell r5 open (e.g., Figs 141, 193, 195, 229)...... 125 125. Female ovipositor at least 2 × as long as high (in lateral view) (as in Figs 252, 253)...... 126 – Female ovipositor < 2 × as long as high (in lateral view) (Figs 254, 255) ...... 130 126. Hind coxa with at least one lateral macroseta together with fine setae (e.g., Figs 17, 230)...... 127 – Hind coxa with fine setae only (without macrosetae)...... 129 127. Discal wing cell (d) markedly constricted at mid length (Fig. 229) ...... Synolcus Loew – Discal wing cell (d) not markedly constricted at mid length (e.g., Fig. 141)...... 128 128. Head with facial swelling smoothly (if only slightly) convex (Fig. 142); antennal stylus as long as or slight- ly longer than postpedicel (Fig. 142); thorax with scutum not markedly humped (Fig. 142); scutal mane (i.e., long, tightly-packed, fine setae arranged in narrow strip mid-dorsally) undeveloped (Fig. 142)...... Dasophrys Loew – Head with facial swelling undeveloped; antennal stylus shorter than postpedicel; thorax with scutum markedly humped; scutal mane well-developed ...... Gibbasilus Londt 129. Scutellum with > 6 apical macrosetae; scutal mane (i.e., long, tightly-packed, fine setae arranged in narrow strip mid-dorsally) well-developed, but without clearly discernible acrostichal macrosetae...... Hippomachus Engel – Scutellum with 6 or fewer apical macrosetae; scutal mane well-developed with clearly discernible acrostichal macrosetae...... Labarus Londt 130. Female terminalia with ovipositor laterally compressed, distinctly longer than high (in lateral view) (Figs 254, 255); cercus smoothly rounded distally (Fig. 255); male terminalia (Figs 240–242), with abdominal sternite 8 with bifurcate medial process distally; phallus long, Z-shaped, each straight section being of similar length and general development ...... Millenarius Londt – Female terminalia with ovipositor usually tubular in form, but if somewhat laterally compressed, then never distinctly longer than high (in lateral view); male terminalia (Figs 243–235), with abdominal stern- ite 8 usually without medial process distally (if process present, then never bifurcate, but knob-like or in form of smoothly-rounded dorsoventrally flattened projection); male phallus short to moderately long, usually fairly straight, but if somewhat Z-shaped, then basal section invariably much more robust than other sections ...... Neolophonotus Engel 131 131. Hind coxa with fine setae only; thorax with postpronotal lobe invariably with setae ...... 132 – Hind coxa with at least 1 (often more) lateral macroseta in addition to fine setae; thorax with postprono- tal lobe with or without setae...... 133 132. Thorax with scutal mane (i.e., long, tightly-packed, fine setae arranged in narrow strip mid-dorsally), usually weakly developed, with only weak, loosely arranged, usually pale coloured setae (rarely absent) in posterior part or absent anteriorly; if scutal mane present, then not bicoloured (black anteriorly, pale yellow or white posteriorly)...... Neolophonotus angustibarbus-group – Thorax with scutal mane well-developed, with longish setae arranged loosely or as tightly-packed row; scutal mane unicolourous black along entire length (may be bordered by smaller pale coloured setae) ...... Neolophonotus suillus-group 133. Thorax with postpronotal lobes setose with several setae...... 134 – Thorax with postpronotal lobes bare (rarely with 1–3 isolated, erect setae)...... 135 134. Thorax with scutal mane (i.e., long, tightly-packed, fine setae arranged in a narrow strip mid-dorsally), weakly to moderately developed, bicolourous (black setae anteriorly, white setae posteriorly) ...... Neolophonotus chionthrix-group – Thorax with scutal mane usually well-developed, unicolourous black (may be bordered by pale colour- ed setae; exceptions with white setae anteriorly and black setae posteriorly) ...... Neolophonotus comatus-group 135. Thorax with scutal mane bicolourous (black setae anteriorly, black or yellow-white setae posteriorly) or unicolourous (white setae along entire length)...... Neolophonotus pellitus-group – Thorax with scutal mane unicolourous black (often bordered by pale coloured setae; exceptions with white setae anteriorly and black setae posteriorly)...... Neolophonotus squamosus-group 136. Scutellum with 4 or more apical macrosetae ...... 137 – Scutellum with 3 or fewer apical macrosetae (e.g., Figs 95, 127)...... 138

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1125

1 137. Head with facial swelling confined to lower ∕2 of face; male terminalia with phallus straight...... Eremisca Hull – Head with facial swelling well-developed; male terminalia with phallus strongly curved ...... Loew 138. Female abdominal sternites 1–5 pubescent, sternites 6–10 shiny and bare; female ovipositor telescopic, comprised of segment 6 and following abdominal segments...... Astochia Becker – Female abdominal sternites 1–6 (at least) pubescent; female ovipositor more or less conical, comprised of segment 8 and following abdominal segments...... 139 139. Antennal stylus composed of 2 articles (1 segment-like article, plus apical seta-like sensory article) (some Malagasy species with 3 articles) ...... 140 – Antennal stylus composed of 3 articles (2 clearly evident segment-like articles, plus apical seta-like sen- sory article) ...... 141 140. Mid femur swollen and with conspicuous cluster of well-developed macrosetae ventrally; proboscis curved upwards distally; thorax with dorsocentral macrosetae developed only postsuturally; female sternites 1–6 pubescent, sternites 7–9 shiny and bare; female cercus spinose, with well-developed macrosetae ...... Hoplopheromerus Becker & Curvirostris Tomasovic – Mid femur not conspicuously swollen, without cluster of well-developed macrosetae ventrally; probos- cis straight; thorax with dorsocentral macrosetae developed pre- and postsuturally; female sternites 1–7 pubescent, sternites 8–9 shiny and bare; female cercus finely setose ...... Heligmonevra Bigot 141. Thorax with postpronotal lobe (and much of scutum) clothed in uniformly short setae (some species have setae of intermediate length, Fig. 96); hind coxa usually with 2 lateral macrosetae (Fig. 96); male terminalia with epandrium with characteristic sub-apical dorsomedial lobe; male phallus straight, later- ally compressed with at most tiny distal prongs; female ovipositor relatively short, only slightly laterally compressed distally...... Afromochtherus Lehr – Thorax with postpronotal lobe with fine, long setae (e.g., Fig. 128); hind coxa usually with 1 lateral mac- roseta (Notomochtherus with ca 3 long, weak setae, barely differentiated from accompanying setae, e.g., Fig. 128); male terminalia with epandrium and phallus differently developed; female ovipositor of various forms ...... 142 142. Male terminalia with phallus strongly curved, exceptionally long, often coiled; terminal abdominal segments with characteristic deep (laterally compressed) appearance; hypandrium not markedly con- stricted medially ...... Valiraptor Londt – Male terminalia with phallus more or less straight to bowed; terminal abdominal segments tubular, not laterally compressed; hypandrium slightly (Notomochtherus) to markedly constricted medially. . . .143 143. Hind femur uniformly dark red-brown to black (proximal or distal end may be paler) (e.g., Fig. 6). . 144 – Hind femur chiefly yellow, with or without distinct dark red-brown or blackish marks or fascia (e.g., Figs 9, 121)...... 146 144. Hind coxa with ca 3 weakly developed lateral macrosetae; male terminalia with phallus abruptly bent upwards at virtually mid length; female ovipositor broader than deep . . . . Notomochtherus Londt – Hind coxa with single lateral macroseta; male terminalia with phallus more of less straight or with slight curve...... 145 145. Thorax with dorsocentral setae well-developed pre- and postsuturally; female terminalia with cercus finely setose; ovipositor laterally compressed; male terminalia with phallus of characteristic shape (shaft gently bowed, prongs short and weakly developed, pump proximally situated (see Theodor 1976))...... Loew – Thorax with dorsocentral setae only well-developed postsuturally (Figs 6, 128); female terminalia with cercus usually spinose, short macrosetae dorsally or finely setose with strongly sclerotised, upturned tip; ovipositor conical; male terminalia with phallus of characteristic development (more or less straight or slightly sinuous, prongs short and moderately well-developed, lateral pair upwardly directed, median prong downwardly directed, sperm pump distally situated (see Londt & Tsacas 1987)) ...... Congomochtherus Oldroyd 146. Hind femur almost entirely yellow (may have poorly defined pale brown patches or dark distal ends) (e.g., Fig. 122)...... 147 – Hind femur with well-defined dark red-brown patches or fascia (other than dark distal ends) (e.g., Fig. 9)...... 150

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1126 SURICATA 5 (2017)

147. Antennal scape and trochanters blackish (e.g., Fig. 122) ...... 148 – Antennal scape and trochanters yellowish ...... 149 148. Confined to Southern Africa...... Caenoura Londt – Confined to Central Africa (T. kivuensis (Tsacas, 1969))...... Tsacasiella Lehr [in part] 1 149. Head with facial swelling occupying ventral ∕2 of face and moderately developed; male terminalia not greatly elongate; epandrium with dorsomedial process; male gonocoxite not markedly elongate; gonostylus elongate, considerably longer than gonocoxite; phallus downwardly-curved distally, prongs minute...... Sphagomyia Londt 1 – Head with facial swelling occupying < ∕2 of face and only weakly developed; male terminalia greatly elongate; epandrium usually with small ventrodistal process; male gonocoxite, gonostylus and phallus long, narrow and relatively straight...... Tsacasiella Lehr [in part] 150. Male abdominal sternite 8 with prominent distomedial process...... Gongromyia Londt – Male abdominal sternite 8 without prominent distomedial process...... 151 151. Antennal postpedicel at least 2 × as long as scape; male terminalia with epandrium characteristically shaped, shortish, converging strongly distally; phallus bent abruptly upwards at virtually mid length and with 3 short, well-developed terminal prongs...... Dikowmyia Londt – Antennal postpedicel < 2 × as long as scape; male terminalia with epandrium and phallus of different shape...... 152 152. Male terminalia with epandrium with complicated arrangement of lobes and processes distally; female ovipositor shortish, broad proximally, laterally compressed distally...... Aneomochtherus Lehr – Male terminalia with epandrium simple, without lobes and processes; female ovipositor longish, later- ally compressed for most of length ...... Melouromyia Londt

Synopsis of the fauna and woodland habitats and adults perch either on the ground or within or at the tips of shrubs and bushes. The oviposition Ecological comments below follow Londt (1994c) and are strategy remains unknown, but probably involves random based on personal observations as well as more recently availa- egg-dropping. Other useful references on the genus include ble field photographs. The oviposition strategy is likewise based Londt (2010b, 2012a) and Tomasovic & De Bakker (2010). on personal observations together with ovipositor morphology. Scarbrough (2005) provided an identification key to species.

Acasilus Londt (Asilinae). An endemic monotypic genus, Afroholopogon Londt (Brachyrhopalinae). An endemic ge- with the single medium-sized species, A. tigrimontis Londt, nus of 19 described species of small- to medium-sized, bee- 2005, confined to South Africa (Londt 2005a). The species in- like species (Figs 18, 19, 91–94) (Londt 2005b). The distri- habits semi- habitat and adults probably perch within bution of the genus is centred in Southern Africa (Botswana, and/or at the tips of shrubs and bushes. Oviposition probably Namibia, South Africa, Swaziland and Zimbabwe), but is also takes place on or in vegetation. recorded from East Africa (Eritrea and Kenya) and the south- ern Arabian Peninsula (Yemen and Oman). The genus inhabits Acnephalomyia Londt (Willistonininae). An endemic genus grass in Fynbos, grassland, savanna, semi-desert and woodland of seven small- to medium-sized, bee-like species (Figs 66, 85– habitats and adults perch either on rocks or within, or at the 88), confined to Southern Africa (Namibia and South Africa) tips of grasses, shrubs and bushes (Figs 18, 19). Oviposition (Londt 2010c). The genus inhabits desert, Fynbos, grassland, takes place in sand or soil. Other useful references to the genus savanna, semi-desert and woodland habitats and adults perch include Londt (1994a). Londt (2005b) provided an identifica- on stones or on the ground (Fig. 66). Oviposition takes place tion key to all species. in sand or soil. Londt (2010c) provided an identification key to the seven species. Afromelittodes Oldroyd & Van Bruggen (Laphriinae). An endemic genus of two large-sized, bee-mimicking species con- Afroestricus Scarbrough (Ommatiinae). An endemic ge- fined to Southern Africa: A. mimos Londt, 2003 and A. solis nus of 20 species of small- to large-sized flies (Figs 89, 90) Oldroyd & Van Bruggen, 1963 (both from Namibia and South described by Scarbrough (2005). The genus is widely distrib- Africa) (Londt 2003). The genus inhabits savanna and wood- uted, in Southern Africa (Botswana, Mozambique, Namibia, land habitats and adults perch within trees. Oviposition takes South Africa and Zimbabwe), Central Africa (Angola, Came- place on or in vegetation. Other useful references to the genus roon, Chad, Democratic Republic of Congo, Gabon, Malawi, include Oldroyd & Van Bruggen (1963). Londt (2003) provid- Republic of Congo and Zambia), East Africa (Burundi, Eritrea, ed an identification key to the two species. Ethiopia, Kenya, Rwanda, Tanzania and Uganda), the southern Arabian Peninsula (Yemen), West Africa (Gambia, Ghana, Mali, Afromochtherus Lehr (Asilinae). An endemic genus of 15 Mauritania, Niger and Nigeria) and the Indian Ocean islands medium- to large-sized species (Figs 95, 96) reviewed by (Madagascar) (Scarbrough 2005). The genus inhabits savanna Londt (2002b) and Londt & Vieira (2013). The genus is fairly

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1127 widespread throughout the Afrotropics and occurs in Southern savanna, semi-desert and woodland habitats and adults perch Africa (Botswana, Namibia, South Africa and Zimbabwe), Cen- on the ground, on stones and rocks, within shrubs and bushes, tral Africa (Democratic Republic of Congo, Malawi and Zam- or at the tips of branches. Oviposition takes place in sand or bia), East Africa (Kenya and Tanzania) and the Indian Ocean soil. Other useful references to the genus include Tomasovic islands (Madagascar). The genus inhabits Fynbos, grassland, (2006) and Tsacas (1969). An identification key to continental

Figs 48.69–74. Heads of Afrotropical Asilidae (anterior views): (69) Afroscleropogon sp. ♀; (70) Corymyia antimelas Londt ♂; (71) Damalis femoralis Ricardo ♂; (72) Dasophrys androclea (Walker) ♂; (73) Daspletis setithoracicus (Ricardo) ♂; (74) Microstylum sp. ♂.

Abbreviations: fr – frons; oc tr – ocellar triangle; tnt pit – tentorial pit; vrt – vertex.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1128 SURICATA 5 (2017)

Afrotropical species was provided by Londt (2002c) and the 2015, confined to Central Africa (Cameroon) (Londt 2015a). Malagasy species by Londt & Vieira (2013). The habitat and of the species remain unknown, but it probably occurs at forest margins or in woodland savanna. Ovi- Afromosia Londt (Laphriinae). An endemic monotypic ge- position probably takes place through random egg-dropping nus, with the single small-sized species, A. barkemeyeri Londt, or into vegetation.

Figs 48.75–80. Heads of Afrotropical Asilidae (anterior views): (75) Neolophonotus sp. ♂; (76) Pegesimallus calvifrons Londt ♀; (77) Promachus amastrus (Walker) ♂; (78) Prytanomyia kochi (Lindner) ♀; (79) Sisyrnodytes subater Oldroyd ♀; (80) Sporado­ thrix gracilis Hermann ♂.

Abbreviations: fr – frons; oc tr – ocellar triangle; tnt pit – tentorial pit; vrt – vertex.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1129

Afroscleropogon Londt (Stenopogoninae). An endemic 1936, 1937, 1938, 1939) and Engel & Cuthbertson (1934). genus of seven medium-sized species (Figs 69, 97, 98) con- Other useful references include Engel & Cuthbertson (1934), fined to Southern Africa (Botswana, Namibia and South Africa) Londt (2005a, 2010b) and Oldroyd (1974). No identification (Londt 1999d). The genus inhabits Fynbos, grassland, savanna key to Afrotropical species is currently available. and semi-desert habitats and adults perch on the ground. Ovi- position takes place in sand or soil. A useful additional refer- Ammodaimon Londt (Willistonininae). A small endemic ence is Oldroyd (1974), who treated species under the name genus of two small-sized, bee-like species (Figs 101, 102), A. Stenopogon Loew. Londt (1999d) provided an identification acares Londt, 1985 and A. platythrix Londt, 2010, confined to key to species. Southern Africa (Namibia) (Londt 1985c). Unpublished records indicate that the genus also occurs in South Africa. The genus Agrostomyia Londt (Brachyrhopalinae). An endemic mono- inhabits grassland and semi-desert habitats and adults perch typic genus, with the single medium-sized species, A. dimor­ on the ground. Oviposition takes place in sand or soil. Londt pha Londt, 1994, confined to Southern Africa (Namibia and (2010c) provided an identification key to the two species. South Africa) (Londt 1994b). Other useful references include Londt (1999d). The species inhabits savanna and grassland Ammophilomima Enderlein (Leptogastrinae). A medium- habitats and adults perch on the ground or within grass. Ovi- sized genus of 41 large-sized species (Figs 103, 104), recorded position takes place in sand or soil. from the Afrotropical and Oriental Regions and reviewed by Janssens (1953) and Martin (1973). The ten Afrotropical spe- Akatiomyia Londt (incerta sedis). An endemic monotypic cies are recorded from Central Africa (Cameroon and Dem- genus, with the single small-sized, black species, A. eremnos ocratic Republic of Congo), East Africa (Burundi and Kenya) Londt, 2013, confined to Southern Africa (South Africa) (Londt and West Africa (Nigeria), although most species are recorded 2013a). The species inhabits Fynbos habitats and adults perch from Democratic Republic of Congo. There are unpublished at the tips of shrubs and bushes. Oviposition takes place in records of the genus from Angola, Central African Republic, sand or soil. Côte d’Ivoire, Equatorial Guinea, Gabon, Ghana, Liberia, Ma- lawi, Mozambique, Togo and Uganda. The biology remains Alcimus Loew (Asilinae). A medium-sized genus of 18 de- unknown, but the genus probably inhabits forest habitats and scribed very large-sized species (Figs 2, 3, 99, 100), requiring adults perch within or at the tips of shrubs and bushes. Ovi- modern revision. Currently, considered an Afrotropical en- position probably takes place through random egg-dropping. demic, although it may occur in the south-western Palaearc- Other useful references include Janssens (1955) and Oldroyd tic Region. The genus is widespread throughout sub-Saharan (1970). Martin (1973) provided an identification key to Afro- Africa, being recorded from Southern Africa (South Africa and tropical species. Zimbabwe), Central Africa (Malawi), East Africa (Ethiopia, Kenya, Sudan, Tanzania and Uganda) and West Africa (Gam- Amphisbetetus Hermann (Brachyrhopalinae). A genus of bia and Senegal). The genus inhabits grassland, savanna and 12 small-sized, often bee-like species (Figs 105, 106), primar- woodland habitats and adults perch on the ground, including ily known from the south-western Palaearctic Region, but also river banks, on rocks, or at the tips of shrubs and bushes (Figs recorded from the Australasian Region. The single Afrotropical 2, 3). Oviposition takes place in sand or soil. Some biological species, A. hermanni Bosák & Hradský, 2011, is confined to information was provided by Cuthbertson (1933, 1934, 1935, the southern Arabian Peninsula (United Arab Emirates) (Bosák

Figs 48.81–82. Heads of Afrotropical Asilidae (anterior views): (81) Stichopogon hermanni Bezzi ♂; (82) Stiphrolamyra diaxantha (Hermann) ♂.

Abbreviations: fr – frons; oc tr – ocellar triangle; vrt – vertex.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1130 SURICATA 5 (2017)

& Hradský 2011: 729). Other useful references include Ef- Oriental and Palaearctic Regions. The 27 Afrotropical species flatoun (1937) and Theodor (1980). The biology remains un- were reviewed by Londt (2011) and are widespread through- known, but the genus probably inhabits semi-desert habitats out sub-Saharan Africa, recorded from Southern Africa (Bot- and adults probably perch on the ground or on low vegetation. swana, Lesotho, Mozambique, Namibia, South Africa, Swa- Oviposition takes place in sand or soil. ziland and Zimbabwe), Central Africa (Angola, Democratic Republic of Congo, Malawi and Zambia), East Africa (Burundi, Anasillomos Londt (Stenopogoninae). An endemic genus of Eritrea, Kenya, Somalia, Sudan and Tanzania) and West Africa two medium-sized species (Figs 107, 108), A. chrysopos Londt, (Côte d’Ivoire, Gambia, Ghana and Nigeria). The genus inhab- 1983 and A. juergeni Dikow, 2015, confined to Southern Af- rica (Botswana, Namibia and South Africa) (Dikow 2015). The its forest (possibly), Fynbos, grassland, savanna, semi-desert genus inhabits grassland, savanna and semi-desert habitats and and woodland habitats and adult perch on the ground, on adults perch on the ground. Oviposition takes place in sand or rocks and at the tips of grass. Oviposition takes place in sand or soil. Other useful references include Dikow & Londt (2000b) soil. Some biological information was provided by Cuthbertson and Londt (1983c). Dikow (2015) provided an identification (1935, 1936). Other useful references include Bromley (1936), key to the two species. Oldroyd (1970, 1974) and Londt (2010b). Londt (2011) pro- vided an identification key to Afrotropical species. Ancylorhynchus Berthold (Stenopogoninae). A large genus of 50 medium- to large-sized, wasp-like species (Figs 109, 110, Andrenosoma Rondani (Laphriinae). A genus of 64 medi- 292–294, 298–300), occurring in the Afrotropical, Australasian, um- to large-sized species, occurring in all zoogeographical

Figs 48.83–84. Wings of Afrotropical Asilidae (dorsal views): (83) Hoplistomerus nobilis Loew; (84) Promachus amastrus (Walker).

Abbreviations: A1 – first branch of anal vein; al – alula; aux vn – auxiliary vein; bm – basal medial cell; br – basal radial cell; C – costal vein; CuA – anterior branch of cubital vein; cua – anterior cubital cell; CuP – posterior branch of cubital vein; cup –

posterior cubital cell; d – discal cell; fal vn – false vein; M1 – first branch of media; m1 – first medial cell; M2 – second branch of media; m2 – second medial cell; M3 – third branch of media; m3 – third medial cell; M4 – fourth branch of media; m4 – fourth medial cell; m–cu – medial–cubital crossvein; m–m – medial crossvein; R1 – anterior branch of radius; r1 – first radial cell; R2+3 – second branch of radius; r2+3 – second + third radial cell; r4 – fourth radial cell; R4 – upper branch of third branch of radius; R4+5 – third branch of radius; r5 – fifth radial cell; R5 – lower branch of third branch of radius; R5+M1 – lower branch of third branch of radius + first branch of media; r–m – radial–medial crossvein; Sc – subcostal vein.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1131 regions, except Antarctica. Two species occur in the Afro- adults probably perch within shrubs, bushes and trees. Ovi- tropical Region, A. boranicum Corti, 1895 and A. serpentina position takes place in vegetation. The immature stages of the (Bezzi, 1908), that are widespread throughout sub-Saharan extralimital species, Andrenosoma cruentum (McAtee, 1919) Africa, recorded from Southern Africa (South Africa and (Figs 320–323) were described by Dennis & Barnes (2014). Zimbabwe), Central Africa (Democratic Republic of Congo Other useful references include Engel & Cuthbertson (1937), and Malawi), East Africa (Ethiopia and Kenya) and West Afri- Oldroyd (1970), the unpublished thesis of Fisher (1986) and ca (Côte d’Ivoire). The biology is poorly known, but species Tomasovic (2010). No identification key to Afrotropical spe- probably inhabit forest, savanna and woodland habitats and cies is currently available.

Figs 48.85–90. Habitus photographs of Asilidae: (85) Acnephalomyia sp. ♂, dorsal view; (86) same, lateral view; (87) Acnephalo­ myia sp. ♀, dorsal view; (88) same, lateral view; (89) Afroestricus chiastoneurus (Speiser) ♂, dorsal view; (90) same, lateral view.

Abbreviations: C – costal vein; CuA+CuP – anterior branch of cubital vein + posterior branch of cubital vein; dc s – dorsocentral

seta; fc – face; m3 – third medial cell; myst – mystax; pped – postpedicel; prbs – proboscis; R4 – upper branch of third branch of radius; r5 – fifth radial cell; scp – scape; sctl – scutellum; st vn – stump vein; styl – stylus.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1132 SURICATA 5 (2017)

Aneomochtherus Lehr (Asilinae). A genus of 63 medium- perch within and/or at the tips of trees and shrubs. Oviposition sized species, occurring in the Afrotropical, Oriental and takes place on or in vegetation. Londt (2002b) provided an Palaearctic Regions. The three Afrotropical species, A. afri­ identification key to Afrotropical species. canus (Ricardo, 1919), A. deserticolus (Karsch, 1888) and A. monobia (Speiser, 1910), are confined to East Africa (Kenya Antiscylaticus Londt (Brachyrhopalinae). An endemic and Tanzania) and were reviewed by Londt (2002b). Habitat monotypic genus, with the single medium-sized species, A. requirements of the genus remain unknown, but probably snowi Londt, 2010, confined to West Africa (Gambia) (Londt comprise savanna and woodland habitats and adults probably 2010b, 2012a). The species inhabits grassland habitats and

Figs 48.91–96. Habitus photographs of Asilidae: (91) Afroholopogon sp., dorsal view ♂; (92) same, lateral view; (93) Afroholopo­ gon sp., dorsal view ♀; (94) same, lateral view; (95) Afromochtherus mendax (Tsacas), dorsal view ♂; (96) same, lateral view.

Abbreviations: anatg – anatergite; C – costal vein; CuP – posterior branch of cubital vein; cup – posterior cubital cell; cx – coxa;

myst – mystax; pprn lb – postpronotal lobe; r1 – first radial cell; R2+3 – second branch of radius; sctl – scutellum.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1133 adults probably perch on the ground or within and at the tips savanna, semi-desert and woodland habitats and adults perch of grass. Oviposition takes place in sand or soil. on the ground, but have also been observed to perch on low vegetation (Figs 25, 26). The oviposition strategy is unknown, Anypodetus Hermann (Laphriinae). An endemic genus of but probably takes place directly into sand. Other useful ref- eight medium-sized, bee-like species (Figs 25, 26, 111, 112, erences include Oldroyd (1974). Londt (2000a) provided an 264–266) reviewed by Londt (2000a). The genus is more or identification key to species. less confined to Southern Africa (Botswana, Mozambique, Na- mibia, South Africa and Zimbabwe) and Central Africa (Zam- Apoclea Macquart (Asilinae). A genus of 21 medium-sized bia). The genus inhabits open sandy places in forest, grassland, species (Figs 4, 5, 113–116), occurring in the Afrotropical,

Figs 48.97–102. Habitus photographs of Asilidae: (97) Afroscleropogon sp., dorsal view ♀; (98) same, lateral view; (99) Alcimus sp., dorsal view ♂; (100) same, lateral view; (101) Ammodaimon sp., dorsal view ♂; (102) same, lateral view.

Abbreviations: anatg – anatergite; aux vn – auxiliary vein; dc s – dorsocentral seta; fc – face; ktg – katatergite; R2+3 – second branch of radius; r4 – fourth radial cell; R4 – upper branch of third branch of radius; R4+5 – third branch of radius; sctl – scute- llum; styl – stylus.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1134 SURICATA 5 (2017)

Oriental and Palaearctic Regions. Four species are recorded (2011), Efflatoun (1934, 1937), Londt (2005a) and Walker & from the Afrotropics, from East Africa (Sudan), the southern Pittaway (1987). No identification key to Afrotropical species Arabian Peninsula (Oman, United Arab Emirates and Yem- is currently available. en) and West Africa (Niger). Unpublished records are also known for Djibouti and Senegal. The genus inhabits desert Astiptomyia Londt (Willistonininae). An endemic monotyp- and semi-desert habitats and adults perch on the ground or on ic genus, with the single small, bee-like species, A. bikos Londt, low vegetation (Figs 4, 5). Oviposition probably takes place in 2010 (Figs 117, 118), confined to Southern Africa (Namibia) sand or soil. Other useful references include Bosák & Hradský (Londt 2010c). The species inhabits desert and semi-desert

Figs 48.103–108. Habitus photographs of Asilidae: (103) Ammophilomima auripennis Janssens, dorsal view ♀; (104) same, lateral view; (105) Amphisbetetus favillaceus (Loew), dorsal view ♂ (non-Afrotropical); (106) same, lateral view; (107) Anasillomos juergeni Dikow, dorsal view ♀; (108) same, lateral view.

Abbreviations: anatg – anatergite; aprn – antepronotum; C – costal vein; CuP – posterior branch of cubital vein; cup – posterior

cubital cell; fem – femur; myst – mystax; ocp – occiput; pal s – postalar seta; prbs – proboscis; prepst – proepisternum; r5 – fifth radial cell; sct – scutum; styl – stylus; tb– tibia.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1135 habitats and adults probably perch on the ground. Oviposition African species, while Londt (2015a) discussed the genus and takes place in sand or soil. transferred species then catalogued as to this and oth- er genera. Species of the genus are widespread throughout Astochia Becker (Asilinae). A genus of 44 large-sized spe- sub-Saharan Africa, recorded from Southern Africa (Mozam- cies, occurring in the Afrotropical, Australasian (possibly), bique, South Africa and Zimbabwe), Central Africa (Came- Oriental and Palaearctic Regions (Geller-Grimm 2004). Five roon, Democratic Republic of Congo, Gabon, Malawi and Re- species reviewed by Londt (1982b) occur in the Afrotropics public of Congo), East Africa (Uganda) and West Africa (Côte that are widespread throughout sub-Saharan Africa, recorded d’Ivoire, Gambia, Ghana, Liberia, Nigeria and Sierra Leone). A from Southern Africa (South Africa and Zimbabwe), Central single record for an Indian Ocean island (Seychelles) requires Africa (Democratic Republic of Congo and Malawi), East Africa confirmation. The genus inhabits forest habitats (including (Kenya, Tanzania and Uganda), the southern Arabian Peninsula thick coastal bush) and adults perch at the tips of shrubs, bush- (Yemen) and West Africa (Nigeria and Sierra Leone). The genus es and trees and within trees. Oviposition takes place on or inhabits savanna and woodland habitats and adults perch at in vegetation (wood). Some biological information (as Laphria) the tips of grass, shrubs and bushes. Oviposition takes place was provided by Cuthbertson (1936). Other useful references on or in vegetation. Other useful references include Londt include Oldroyd (1970), who produced a key to Afrotropical (2002b). Londt (1982b) provided an identification key to the Laphria, that includes species now assigned to Choerades, Ol- five Afrotropical species. droyd (1974), who keyed the Southern African species (also under the name Laphria), Bromley (1935), Tomasovic (2007, Bana Londt (Stenopogoninae). An endemic genus described by 2008a) and Londt (2015a). No useful identification key to Londt (1992b), comprising two large-sized, bee-like species (Figs Afrotropical species is currently available. 119, 120), B. apicida Londt, 1992 and B. madiba Londt, 2013, confined to Southern Africa (Namibia and South Africa) (Londt Clinopogon Bezzi (Stichopogoninae). A genus of nine 2013b). The genus inhabits grassland, savanna and semi-desert medium-sized species (Figs 125, 126), occurring in the Afro- habitats and adults perch on the ground or at the tips of grass, tropical, Australasian, Oriental and Palaearctic Regions. Two shrubs and bushes. Oviposition takes place in sand or soil. Londt species occur in the Afrotropics, C. nicobarensis (Schiner, (2013b) provided an identification key to the two species. 1868) and C. reginaldi (Séguy, 1955). This littoral genus is dis- tributed along almost the entire eastern coastline of Africa Caenoura Londt (Asilinae). An endemic genus of two small- (C. nicobarensis) and Tromelin Is. (C. reginaldi). The genus to medium-sized species (Figs 121, 122), described by Londt inhabits maritime beach habitats and adults perch on the (2002b), C. annulitarsis (Loew, 1858) and C. sinuatus (Loew, ground. Oviposition takes place in sand or soil. Other useful 1858), confined to Southern Africa (Lesotho, South Africa and Zimbabwe). The genus inhabits grassland, savanna and wood- references include Londt & Copeland (2017), Oldroyd (1974) land habitats and adults perch within grass. Oviposition takes and Séguy (1955). Although Londt (1979b) briefly reviewed place on or in vegetation. Londt (2002b) provided an identifi- the Afrotropical species, the genus requires modern revision cation key to the two species. and no identification key to Afrotropical species is currently available. Caroncoma Londt (Dasypogoninae). An endemic monotyp- ic genus described by Londt (1980b), with the single medium- Congomochtherus Oldroyd (Asilinae). An endemic genus sized species, C. atrimaculatus (Oldroyd, 1960), confined to of seven medium-sized species (Figs 6, 127, 128), reviewed the Indian Ocean islands (Madagascar). The species proba- by Londt & Tsacas (1987) and Londt (2014c). The genus is bly inhabits forest habitats and adults probably perch on the widespread throughout sub-Saharan Africa, recorded from ground or within or at the tips of shrubs and bushes. Oviposi- Southern Africa (Namibia, South Africa and Zimbabwe), Cen- tion takes place in sand or soil. Other useful references include tral Africa (Angola, Cameroon, Democratic Republic of Congo, Oldroyd (1960a). Republic of Congo and Zambia), East Africa (Kenya, Tanzania and Uganda) and West Africa (Nigeria). The genus inhabits Cerdistus Loew (Asilinae). A genus of 64 medium-sized spe- Fynbos, grassland, savanna and woodland habitats and adults cies, occurring in the Afrotropical, Australasian (doubtfully), perch on the ground, on stones, riverbanks and other riparian Oriental and Palaearctic Regions (Geller-Grimm 2004). The habitats, including rocks surrounded by water (Fig. 6). Ovipo- single Afrotropical species, C. griseola (Oldroyd, 1960), was sition probably takes place in riverine sand or soil. Other useful recorded from Madagascar by Oldroyd (1960b), but remains references include Londt (2002b) and Oldroyd (1970). Londt doubtful and requires revision, along with Palaearctic species. (2014c) provided an identification key to the seven species. Geller-Grimm (2002) further records an undescribed species from Socotra Is. (Yemen). The genus inhabits either forest Connomyia Londt (Stenopogoninae). A near endemic genus (Madagascar) or semi-desert (Socotra Is.) habitats and adults of 22 medium-sized, bee-like species (Figs 129, 130), occurring perch within and/or at the tips of shrubs and bushes. Ovipo- in the Afrotropical and Oriental Regions. The 20 Afrotropical sition probably takes place on or in vegetation. Other useful species were reviewed by Londt (1993c) and are widespread references include Londt (2002b). throughout sub-Saharan Africa, recorded from Southern Afri- ca (Botswana, Lesotho, Mozambique, Namibia, South Africa Choerades Walker (Laphriinae). A genus of 75 medium- and Zimbabwe), Central Africa (Angola, Democratic Republic sized, bee-like species (Figs 27, 123, 124), occurring in the of Congo and Malawi), East Africa (Eritrea, Kenya and Tanza- Afrotropical, Australasian, Oriental and Palaearctic Regions. nia) and West Africa (Burkina Faso, Gambia, Mali, Nigeria and Twenty-five species occur in the Afrotropics, but the genus re- Senegal). The genus inhabits Fynbos, grassland, savanna and quires modern revision. Londt (1977) reviewed the Southern woodland habitats and adults perch on the ground. Oviposition

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1136 SURICATA 5 (2017) takes place in sand or soil. Other useful references include islands of Comoros, Europa Is., Madagascar (incl. Sainte- Londt (2010b). Londt (1993c) provided an identification key Marie Is.), Mauritius (Maurice Is.), Réunion Is., Seychelles (incl. to Afrotropical species. Aldabra and Cosmoledo Is.) and Socotra Is. (Yemen). The bi- ology remains unknown, but species apparently inhabit “dry Cophinopoda Hull (Ommatiinae). A genus of 11 large- to forest, scrub” and other island vegetation and adults probably very large-sized species (Figs 46, 131, 132), occurring in the perch at the tips of shrubs and bushes (Fig. 46). The oviposition Afrotropical, Australasian, Oriental and Palaearctic Regions. strategy probably involves random egg-dropping. Other use- Five species occur in the Afrotropics, that were reviewed by ful references include Martin (1964). Tsacas & Artigas (1994) Tsacas & Artigas (1994) and are confined to the Indian Ocean provided an identification key to the five Afrotropical species.

Figs 48.109–114. Habitus photographs of Asilidae: (109) Ancylorhynchus unifasciatus (Loew), dorsal view ♂; (110) same, lateral view; (111) Anypodetus fascipennis Engel, dorsal view ♂; (112) same, lateral view; (113) Apoclea sp., dorsal view ♂; (114) same, lateral view.

Abbreviations: anatg – anatergite; anepst – anepisternum; dc s – dorsocentral seta; fc – face; pgn – postgena; prbs – proboscis;

R4 – upper branch of third branch of radius; r5 – fifth radial cell; sctl – scutellum; st vn – stump vein.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1137

Corymyia Londt (Stenopogoninae). An endemic genus of Regions. Two species occur in the Afrotropics, C. molitrix Loew, four small-sized species (Figs 70, 133–136), confined to South- 1873 and C. ruficornis (Wulp, 1899), confined to the southern ern Africa (South Africa) (Londt 1994b). The genus inhabits Arabian Peninsula (United Arab Emirates and Yemen) and East grassland, Fynbos and semi-desert habitats and adults perch Africa (Sudan). The genus most probably inhabits semi-desert on the ground. Oviposition takes place in sand or soil. Londt habitats and adults probably perch on the ground, on stones (1994b) provided an identification key to the four species. or in riparian habitats. Oviposition takes place in sand or soil. Other useful references include Efflatoun (1934), who provid- Ctenota Loew (Laphriinae). A genus of seven medium- to ed an excellent description of the type species (C. molitrix), large-sized species, occurring in the Afrotropical and Palaearctic which occurs in Egypt, Hull (1962), Theodor (1980) and Bosák

Figs 48.115–120. Habitus photographs of Asilidae: (115) Apoclea sp., dorsal view ♀; (116) same, lateral view; (117) Astiptomyia bikos Londt, dorsal view ♂; (118) same, lateral view; (119) Bana apicida Londt, dorsal view ♂; (120) same, lateral view.

Abbreviations: anatg – anatergite; anepm – anepimeron; C – costal vein; CuP – posterior branch of cubital vein; cup – posterior

cubital cell; m3 – third medial cell; plp – palpus; R4 – upper branch of third branch of radius; r5 – fifth radial cell; sctl – scutellum.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1138 SURICATA 5 (2017)

& Hradský (2011). No identification key to Afrotropical species Hoplopheromerus key out together in the above identification is currently available. key and these genera are potentially synonymous.

Curvirostris Tomasovic (Asilinae). An endemic monotypic Damalis F. (Trigonomiminae). A large genus of 107 medium- genus, with the single large-sized species, C. dupontae Tomas- sized species (Figs 62, 63, 71, 137–140, 307–309), occurring ovic, 2015, confined to West Africa (Côte d’Ivoire) (Tomas- in the Afrotropical, Oriental and Palaearctic Regions. Thirty- ovic 2015). The species inhabits forest margin habitats and five species are recorded from the Afrotropics, which were adults probably perch at the tips of shrubs, bushes and trees. reviewed by Londt (1989a) and Oldroyd (1960a). The ge- The oviposition strategy remains unknown. Curvirostris and nus is widespread throughout the Afrotropics, recorded from

Figs 48.121–126. Habitus photographs of Asilidae: (121) Caenoura annulitarsis (Loew), dorsal view ♂; (122) same, lateral view; (123) Choerades bella (Loew), dorsal view ♂; (124) same, lateral view; (125) Clinopogon nicobarensis (Schiner), dorsal view ♂; (126) same, lateral view.

Abbreviations: fem – femur; m3 – third medial cell; ped – pedicel; pped – postpedicel; prbs – proboscis; scp – scape; trc – tro- chanter.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1139

Southern Africa (Mozambique, South Africa and Zimbabwe), Dichaetothyrea De Meijere (Laphriinae). A genus of two Central Africa (Angola, Cameroon, Central African Republic, small-sized species, occurring in the Afrotropical and Oriental Democratic Republic of Congo, Malawi and Zambia), East Af- Regions. The single Afrotropical species, D. calvifrons Londt, rica (Kenya, Tanzania and Uganda), West Africa (Benin, Côte 1982, is confined to the Indian Ocean islands (Madagascar) d’Ivoire, Ghana, Guinea, Liberia, Nigeria and Sierra Leone) (Londt 1982a). The biology remains unknown, but the species and the Indian Ocean islands (Madagascar). Aspects of the bi- probably inhabits forest habitats and adults probably perch ology of D. femoralis Ricardo, 1925 were published by Londt within and/or at the tips of shrubs and bushes. Oviposition (1991). The genus inhabits forest (margins), Fynbos, grassland, probably takes place through random egg-dropping. savanna and woodland habitats and adults perch within grass and at the tips of grass, shrubs, bushes and trees (Figs 62, 63). Dichropogon Bezzi (Stichopogoninae). A genus of five Oviposition takes place through random egg-dropping. Some small-sized species, occurring in the Afrotropical and Palaearc- biological information was provided by Cuthbertson (1938, tic Regions. Dichropogon was formerly treated as a subge- as Lophurodamalis Hermann) and eggs and larvae were de- nus of Stichopogon (Geller-Grimm 2004; Hull 1962), but scribed by Londt (1991). Londt (1989a) and Oldroyd (1960a) was recently elevated to full generic status for south-western provided identification keys to the continental Afrotropical and Palaearctic species. A single species, D. tenebrosus Bosák & Malagasy species, respectively. Hradský, 2011, is recorded from the Afrotropics, confined to the southern Arabian Peninsula (United Arab Emirates) (Bosák Danomyia Londt (Stenopogoninae). An endemic genus of & Hradský 2011: 741). The biology of the species remains un- nine medium-sized, bee-like species, centred in Southern Af- known, but is probably similar to that of semi-desert inhabit- rica (Botswana, South Africa and Zimbabwe), but also record- ing Stichopogon species, with adults probably perching on the ed from Central Africa (Chad) and East Africa (Sudan) (Londt ground. Oviposition takes place in sand or soil. 1993c). The genus inhabits grassland, savanna and woodland habitats and adults perch on the ground. Oviposition takes Dikowmyia Londt (Asilinae). An endemic monotypic ge- place in sand or soil. Londt (1993c) provided an identification nus, with the single medium-sized species, D. mediorus key to the nine species. Londt, 2002, confined to Southern Africa (South Africa) (Londt 2002b). The species inhabits grassland habitats and adults Dasophrys Loew (Asilinae). A large endemic genus of 32 probably perch within and at the tips of grass. Oviposition medium- to large-sized species (Figs 7, 72, 141, 142), re- probably takes place in or on vegetation. viewed by Londt (1981b) who later added an additional spe- Dioctobroma Hull (Stenopogoninae). An endemic mono- cies (Londt 1985a). The genus is confined to Southern Africa typic genus, with the single medium-sized species, D. flavoter­ (Lesotho, Mozambique, Namibia, South Africa, Swaziland and minatus Hull, 1962, confined to Southern Africa (Botswana Zimbabwe) and inhabits forest and woodland habitats and and Namibia) (Hull 1962). The biology of the species remains adults perch within and at the tips of shrubs and bushes (Fig. unknown, but it probably inhabits grassland, savanna and 7). Oviposition takes place on or in woody vegetation. Some semi-desert habitats and adults probably perch on the ground. biological information was provided by Cuthbertson (1936). Oviposition takes place in sand or soil. Other useful references Other useful references include Londt (2005a). Londt (1981b) include Londt (1983c) and Dikow & Londt (2000b). provided the most recent identification key to species, but see Londt (1985a) for an additional species. Dogonia Oldroyd (Stenopogoninae). An endemic mono- typic genus, with the single medium- to large-sized species, D. Daspletis Loew (Stenopogoninae). An endemic genus of saegeri Oldroyd, 1970, confined to Central Africa (Democrat- nine medium- to very large-sized species (Figs 49–51, 73, 143, ic Republic of Congo). The genus was described by Oldroyd 144), confined to Southern Africa (Botswana, Namibia, South (1970) and was reviewed by Londt (2008a). The biology of Africa, Zimbabwe) (Londt 2010a), with unpublished records the species remains unknown, but it probably inhabits forest from Mozambique. Tomasovic (2016) described a new species habitats and adults probably perch on the ground or at the tips from Gabon, but the generic assignment is questionable. The of shrubs and bushes. Oviposition takes place in sand or soil. genus inhabits grassland, savanna and semi-desert habitats and adults perch on the ground or on low vegetation (Figs 49–51). Dysclytus Loew (Asilinae). An endemic monotypic genus, Oviposition takes place in sand or soil. Other useful referenc- with the single large-sized species, D. firmatus Walker, 1857, es include Londt (1983c, 1985a) and Tomasovic & Constant confined to Southern Africa (South Africa) (Londt 1979a). The (2013). Londt (2010a) provided the most recent identification species inhabits savanna and woodland habitats and adults key to species, but see Tomasovic (2016) for an additional spe- perch at the tips of shrubs and bushes. Oviposition takes place cies. on or in vegetation. Other useful references include Londt (2005a) and Oldroyd (1974). Dasyllina Bromley (Laphriinae). An endemic monotypic ge- nus, with the single large-sized, bee-like species, D. fulvithorax Emphysomera Schiner (Ommatiinae). A genus of 21 Bromley, 1935, confined to Central Africa (Democratic Repub- medium-sized species (Figs 145, 146), occurring in the Afro- lic of Congo). The biology remains unknown, but the species tropical, Australasian and Oriental Regions. Four species are re- probably inhabits forest or woodland habitats and adults prob- corded from the Afrotropics, that are widespread and recorded ably perch within or at the tips of shrubs, bushes and trees. from Southern Africa (Botswana, Mozambique, South Africa, Oviposition probably takes place on or in vegetation. Other Swaziland and Zimbabwe), Central Africa (Chad, Democratic useful references include Bromley (1935), Londt (1988b) and Republic of Congo, Gabon, Malawi and Zambia), East Afri- Oldroyd (1970). ca (Kenya, Tanzania and Uganda), West Africa (Côte d’Ivoire,

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1140 SURICATA 5 (2017)

Gambia, Ghana, Guinea, Liberia, Senegal and Sierra Leone) Empodiodes Oldroyd (Stenopogoninae). An endemic genus and the Indian Ocean islands (Madagascar) (Scarbrough & of four small- to medium-sized species, confined to Southern Marascia 1996). The genus inhabits forest and woodland hab- Africa (Namibia and South Africa). The genus was described by itats and adults perch at the tips of shrubs and bushes. Ovipo- Oldroyd (1972) and was reviewed by Londt (1992a, 2012c). The sition takes place through random egg-dropping. Other use- genus inhabits grassland, Fynbos and savanna habitats and adults ful references include Londt (2010b). Scarbrough & Marascia perch on the ground. Oviposition takes place in sand or soil. (1996) provided an identification key to the four species. Londt (2012c) provided an identification key to the four species.

Figs 48.127–132. Habitus photographs of Asilidae: (127) Congomochtherus elferinki Londt & Tsacas, dorsal view ♂; (128) same, lateral view; (129) Connomyia leonina (Engel), dorsal view ♂; (130) same, lateral view; (131) Cophinopoda pulchripes (Bigot), dorsal view ♀; (132) same, lateral view.

Abbreviations: anatg – anatergite; anepm – anepimeron; cx – coxa; dc s – dorsocentral seta; myst – mystax; plp – palpus; pprn lb – postpronotal lobe; sctl – scutellum; styl – stylus; term – terminalia.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1141

Erax Scopoli (Asilinae). A genus of 27 medium-sized spe- Londt (2005a). No identification key to Afrotropical species is cies occurring in the Afrotropical and Palaearctic Regions. currently available. Three species, E. albiceps Macquart, 1850, E. costalis (Wulp, 1899) and E. nigrotinctus (Becker, 1909), occur in the Afro- Eremisca Hull (Asilinae). A genus of 17 medium-sized tropics, confined to East Africa (Kenya) and the southern Ara- species (Fig. 8), occurring in the Afrotropical and Palaearctic bian Peninsula (Yemen). The genus probably inhabits desert Regions. The single Afrotropical species, E. heleni (Efflatoun, and semi-desert habitats and adults probably perch on the 1934), is confined to the southern Arabian Peninsula (Oman ground. The oviposition strategy remains unknown, but may and United Arab Emirates) (Lehr 1988b). The genus probably take place in sand or soil. Other useful references include inhabits desert and semi-desert habitats and adults perch on

Figs 48.133–138. Habitus photographs of Asilidae: (133) Corymyia melas Londt, dorsal view ♂; (134) same, lateral view; (135) C. antimelas Londt, dorsal view ♀; (136) same, lateral view; (137) Damalis femoralis Ricardo, dorsal view ♂; (138) same, lateral view.

Abbreviations: acanth sp – acanthophorite spine; anepm – anepimeron; m3 – third medial cell; myst – mystax; plp – palpus; pped – postpedicel; prbs – proboscis; term – terminalia; tg – tergite.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1142 SURICATA 5 (2017) the ground (Fig. 8). Oviposition probably takes place in sand or remains unknown, but it probably inhabits forest habitats and soil. Other useful references include Bosák & Hradský (2011) adults probably perch at the tips of shrubs, bushes and trees. and Theodor (1980). Oviposition may involve random egg-dropping or placement of eggs on or in dead wood. Other useful references include Ericomyia Londt (Laphriinae). An endemic monotypic ge- Oldroyd (1960a). nus, with the single medium-sized, bee-like species, E. ato­ mentosa (Oldroyd, 1960), confined to the Indian Ocean is- Euscelidia Westwood (Leptogastrinae). A large genus of 68 lands (Madagascar) (Londt 2015a). The biology of the species medium-sized species (Figs 42, 147, 148, 275–277), occurring

Figs 48.139–144. Habitus photographs of Asilidae: (139) Damalis annulata Loew, dorsal view ♂; (140) same, lateral view; (141) Dasophrys androclea (Walker), dorsal view ♂; (142) same, lateral view; (143) Daspletis setithoracicus (Ricardo), dorsal view ♂; (144) same, lateral view.

Abbreviations: anatg – anatergite; anepst – anepisternum; d – discal cell; dc s – dorsocentral seta; fc – face; M1 – first branch of media; myst – mystax; pmtcx memb – postmetacoxal membrane; pocl s – postocular seta; pped – postpedicel; r5 – fifth radial cell; sct mn – scutal mane; styl – stylus.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1143 in the Afrotropical, Oriental and Palaearctic Regions. The 55 Sierra Leone and Togo) and the Indian Ocean islands (Mad- Afrotropical species are widespread throughout the region, agascar) (Dikow 2003). The genus inhabits grassland, Fyn- reported from Southern Africa (Botswana, Mozambique, bos, savanna, semi-desert and woodland habitats and adults Namibia, South Africa, Swaziland and Zimbabwe), Central perch at the tips of grass, shrubs, bushes and trees (Fig. 42). Africa (Angola, Cameroon, Central African Republic, Chad, Oviposition takes place through random egg-dropping. Some Democratic Republic of Congo, Malawi and Zambia), East biological information was provided by Cuthbertson (1939). Africa (Burundi, Eritrea, Ethiopia, Kenya, Rwanda, Sudan, Other useful references include Londt (2010b, 2012a). Tanzania and Uganda), West Africa (Burkina Faso, Côte d’Ivo- Dikow (2003) provided an identification key to the 55 Afro- ire, Gambia, Ghana, Guinea, Liberia, Mali, Nigeria, Senegal, tropical species.

Figs 48.145–150. Habitus photographs of Asilidae: (145) Emphysomera pallidapex (Bigot), dorsal view ♂; (146) same, lateral view; (147) Euscelidia procula (Walker), dorsal view ♂; (148) same, lateral view; (149) Gonioscelis sp., dorsal view ♂; (150) same, lateral view.

Abbreviations: al – alula; C – costal vein; fc – face; fem – femur; m3 – third medial cell; pprn – postpronotum; r1 – first radial cell; r4 – fourth radial cell; r5 – fifth radial cell; sctl – scutellum; styl – stylus; tg – tergite.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1144 SURICATA 5 (2017)

Fishermyia Londt (Stenopogoninae). An endemic monotyp- from the Afrotropics, with a disjunct distribution in sub-Saharan ic genus, with the single large-sized species, F. stuckenbergi Africa, recorded from Southern Africa (Namibia and South Af- Londt, 2012, confined to the Indian Ocean islands (Madagas- rica), the southern Arabian Peninsula (United Arab Emirates car) (Londt 2012b). The species inhabits arid spiny woodland and Yemen) and West Africa (Gambia) (Londt 2000b). The ge- habitats and adults perch on the ground. Oviposition takes nus inhabits grassland and savanna habitats and adults perch place in sand or soil. on the ground or on low vegetation (Fig. 20). Oviposition takes place in sand or soil. Other useful references include Bosák & Gerrolasius Hermann (Laphriinae). An endemic genus of Hradský (2011), Kirk-Spriggs & McGregor (2009) and Londt three small species, G. hermanni Londt, 1988, G. meridiona­ (1981a, 1999a, 2000b, 2010b, 2012a). Londt (2000b) provid- lis Hermann, 1920 and G. oldroydi Londt, 1988, confined to ed the most recent identification key to Afrotropical species, Southern Africa (Botswana, Mozambique and South Africa) and but see Bosák & Hradský (2011: 731) for two additional spe- East Africa (Somalia) (Londt 1988b). The genus inhabits grass- cies. land and savanna habitats and adults perch at the tips of grass. Oviposition takes place through random egg-dropping. Londt Haroldia Londt (Stenopogoninae). An endemic genus of two (1988b) provided an identification key to the three species. small species (Figs 153, 154), H. oldroydi Londt, 1999 and H. trivialis (Oldroyd, 1974), confined to Southern Africa (South Af- Gibbasilus Londt (Asilinae). An endemic genus of six rica) (Londt 1999d). The genus inhabits Fynbos and Succulent- medium-sized species confined to Southern Africa (South Af- Karoo habitats and adults perch on the ground (coastal sand rica). The genus was described by Londt (1986a) and was later dunes and beaches). Oviposition takes place in sand or reviewed by Londt (1990b, 2016). The genus inhabits Fynbos soil. Other useful references include Oldroyd (1974). Londt habitats and adults perch at the tips of shrubs and bushes. Ovi- (1999d) provided an identification key to the two species. position takes place on or in vegetation. Londt (2016) provided an identification key to the six species. Heligmonevra Bigot (Asilinae). A genus of 61 medium-sized species occurring in the Afrotropical, Australasian and Orien- Goneccalypsis Hermann (Laphriinae). A genus of four tal Regions. Twenty-four species are recorded from the Afro- small-sized species (Fig. 28) occurring in the Afrotropical and tropics, being widespread throughout the region, including Oriental Regions. Two species occur in the Afrotropics, G. ar­ Southern Africa (Namibia), Central Africa (Bioko Is. (Equatorial genteoviridis (Hermann, 1907) and G. montanus Londt, 1982, Guinea), Cameroon, Democratic Republic of Congo, Gabon, confined to Southern Africa (Lesotho and South Africa) (Londt Malawi and Republic of Congo), East Africa (Eritrea, Kenya, 1982a). The genus inhabits grassland (incl. montane grassland) Tanzania (incl. Zanzibar Is.)), West Africa (Côte d’Ivoire, Gam- and savanna habitats and adults perch at the tips of grass. Ovi- bia and Ghana) and the Indian Ocean islands (Madagascar and position probably takes place through random egg-dropping. Seychelles). The genus inhabits forest, savanna and woodland Londt (1982a) provided an identification key to the two Afro- habitats and adults perch within grass. Oviposition probably tropical species. takes place on or in vegetation. Martin (1964) reviewed the Malagasy fauna (13 species) and other useful references in- Gongromyia Londt (Asilinae). An endemic monotypic genus, clude Engel (1927), Londt (2002b, 2010b, 2012a) and Tomas- with the single medium-sized species, G. bulla Londt, 2002, ovic & Dekoninck (2014). No identification key to Afrotropical confined to Southern Africa (South Africa) (Londt 2002b). The fauna is currently available. species probably inhabits grassland habitats and adults perch within and at the tips of grass. Oviposition probably takes place Hermannomyia Oldroyd (Brachyrhopalinae). An endemic in or on vegetation. genus of three medium-sized species, H. engeli (Hull, 1962), H. oldroydi Londt, 1981 and H. ukazi Londt & Copeland, Gonioscelis Schiner (Stenopogoninae). An endemic genus 2013, confined to Southern Africa (Lesotho, South Africa and of 40 medium-sized species (Figs 52, 149, 150), widely distrib- Zimbabwe) and East Africa (Kenya). The genus inhabits grass- uted throughout sub-Saharan Africa, recorded from Southern land, savanna and woodland habitats and adults perch on the Africa (Botswana, Lesotho, Mozambique, Namibia, South Afri- ground. Oviposition takes place in sand or soil. The genus was ca and Zimbabwe), Central Africa (Angola, Cameroon, Chad, reviewed by Londt (1981a) and Londt & Copeland (2013) Democratic Republic of Congo, Malawi, Republic of Congo and other useful references include Oldroyd (1974). Londt & and Zambia), East Africa (Burundi, Eritrea, Ethiopia, Kenya, Copeland (2013) provided an identification key to the three Tanzania and Uganda) and West Africa (Gambia and Ghana) species. (Londt 2004c). The genus inhabits grassland, Fynbos, savan- na and woodland habitats and adults perch on the ground Hippomachus Engel (Asilinae). A near endemic genus of or on rocks and at the tips of grass, shrubs and bushes (Fig. ten medium-sized species, occurring in the Afrotropical and 52). Oviposition takes place in sand or soil. Some biological Palaearctic Regions. Nine species occur in the Afrotropics that information was provided by Cuthbertson (1936, 1938, 1939). are widespread in sub-Saharan Africa, recorded from Southern Other useful references include Engel (1926), Londt (2010b), Africa (Namibia, South Africa and Zimbabwe), Central Africa Oldroyd (1970, 1974), Ricardo (1925), Tomasovic (2009) and (Angola and Central African Republic), East Africa (Djibouti Tomasovic & Constant (2013). Londt (2004c) provided an and Kenya) and West Africa (Gambia) (Londt 1983b, 1985a, identification key to the 40 Afrotropical species. 2010b). The genus inhabits savanna and woodland habitats and adults perch within trees. Oviposition takes place on or in Habropogon Loew (Brachyrhopalinae). A genus of 43 small- vegetation. Londt (1983b) provided the most recent identifica- sized species (Figs 20, 151, 152, 256–258), occurring in the tion key to Afrotropical species, but see Londt (1985a, 2010b) Afrotropical and Palaearctic Regions. Ten species are recorded for two additional species.

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Hoplistomerus Macquart (Laphriinae). An endemic genus Africa (Gambia, Ghana, Guinea, Mali, Mauritania, Niger, Nige- of 11 medium- to large-sized species (Figs 29, 83, 155, 156, ria and Senegal) (Londt 2007b). Hoplistomerus nobilis was ac- 267, 268), widely distributed throughout sub-Saharan Africa, cidentally introduced into the Neotropical Region (Costa Rica) recorded from Southern Africa (Botswana, Lesotho, Mozam- (Fisher 2009: 622). The genus mainly inhabits grassland and bique, Namibia, South Africa, Swaziland and Zimbabwe), Cen- savanna habitats and adults perch on the ground or on stones tral Africa (Chad, Democratic Republic of Congo, Malawi and (Fig. 29). Oviposition probably takes place in decomposing veg- Zambia), East Africa (Ethiopia, Kenya, Somalia, Sudan, Tanzania etable matter (including dung). The genus feeds almost exclu- and Uganda), the southern Arabian Peninsula (Yemen) and West sively on dung-frequenting beetles (Coleoptera: Scarabaeidae)

Figs 48.151–156. Habitus photographs of Asilidae: (151) Habropogon capensis Londt, dorsal view ♂; (152) same, lateral view; (153) Haroldia trivialis (Oldroyd), dorsal view ♂; (154) same, lateral view; (155) Hoplistomerus nobilis Loew, dorsal view ♀; (156) same, lateral view.

Abbreviations: – antenna; C – costal vein; CuA+CuP – anterior branch of cubital vein + posterior branch of cubital vein; cup – posterior cubital cell; dc s – dorsocentral seta; fem – femur; myst – mystax; ped – pedicel; pped – postpedicel; prbs – proboscis; scp – scape; tsm – tarsomeres.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1146 SURICATA 5 (2017) and some biological information was provided by Cuthbertson Central Africa (Democratic Republic of Congo) and West Af- (1933, 1936, 1939). Other useful references include Londt rica (Burkina Faso, Côte d’Ivoire, Guinea and Nigeria). The (2010b, 2012a) and Oldroyd (1940, 1970, 1974). An identifi- genus inhabits forest habitats and adults perch within trees. cation key to the 11 species is provided by Londt 2007b. Oviposition takes place on or in vegetation. Curvirostris and Hoplopheromerus key out together in the above identification Hoplopheromerus Becker (Asilinae). A genus of ten large- key and these genera are potentially synonymous, with Hoplo­ sized species, occurring in the Afrotropical, Oriental and pheromerus then having priority. Other useful references in- Palaearctic Regions. Five species occur in the Afrotropics that clude Londt (2002b). Tsacas & Oldroyd (1967) provided the are fairly widespread throughout sub-Saharan Africa, including most recent identification key to the five Afrotropical species.

Figs 48.157–162. Habitus photographs of Asilidae: (157) Hypenetes stigmatias Loew, dorsal view ♂; (158) same, lateral view; (159) Hyperechia nigripennis (Wiedemann), dorsal view ♀; (160) same, lateral view; (161) Lamyra gulo (Loew), dorsal view ♂; (162) same, lateral view.

Abbreviations: C – costal vein; fc – face; fem – femur; M2 – second branch of media; mtkepst – metakatepisternum; ovp – ovi- positor; pgn – postgena; plp – palpus; pped – postpedicel; r1 – first radial cell; r5 – fifth radial cell; tb – tibia; tg – tergite.

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Hynirhynchus Lindner (Brachyrhopalinae). An endemic (2010b). Londt (1992a) provided an identification key to the genus of two small- to medium-sized species, H. pantherinus two species. (Bigot, 1879) and H. zebra Lindner, 1955, recorded from East Africa (Kenya and Tanzania) and West Africa (Gambia and Hypenetes Loew (Tillobromatinae). An endemic genus of Senegal) (Londt 1992a). Little is known regarding the biology 21 small- to large-sized species (Figs 61, 157, 158, 303–306), of the genus, but it probably inhabits grassland, savanna and confined to Southern Africa (South Africa) (Londt 1985b). The woodland habitats and adults probably perch on the ground genus inhabits grassland, Fynbos and savanna habitats and or on river banks. Oviposition takes place in sand or soil. adults perch on the ground, on beaches, rocks, or the tips of Other useful references include Lindner (1955) and Londt grass (Fig. 61). Oviposition takes place in sand or soil. Other

Figs 48.163–168. Habitus photographs of Asilidae: (163) Laphyctis sp., dorsal view ♂; (164) same, lateral view; (165) Laphystotes ariel Londt, dorsal view ♂; (166) same, lateral view; (167) Lasiocnemus lugens Loew, dorsal view ♂; (168) same, lateral view.

Abbreviations: anepst – anepisternum; fem – femur; m3 – third medial cell; myst – mystax; ped – pedicel; pgn – postgena; pped – postpedicel; r5 – fifth radial cell; scp – scape; sctl – scutellum; styl – stylus; tb – tibia; term – terminalia.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1148 SURICATA 5 (2017) useful references include Oldroyd (1974). Londt (1985b) pro- South Africa and Zimbabwe), Central Africa (Cameroon, Dem- vided an identification key to the 21 species. ocratic Republic of Congo and Malawi), East Africa (Kenya and Tanzania (incl. Zanzibar)), West Africa (Guinea and Senegal) Hyperechia Schiner (Laphriinae). A genus of 17 large-sized, and the Indian Ocean islands (Madagascar). The genus inhab- (Xylocopa Latreille species; Apidae) mimicking its forest and woodland habitats and adults perch within trees. flies (Figs 159, 160), occurring in the Afrotropical and Oriental Oviposition takes place on or in vegetation (wood). Some bi- Regions, in need of modern revision. Fourteen species occur in ological information was provided by Cuthbertson (1934) and the Afrotropics, which are fairly widespread throughout the re- Poulton (1925) who also illustrated the immature stages. Other gion, recorded from Southern Africa (Mozambique, Namibia, useful references include Enderlein (1930), Grünberg (1907),

Figs 48.169–174. Habitus photographs of Asilidae: (169) Laxenecera mollis (Loew), dorsal view ♀; (170) same, lateral view; (171) Leptogaster aganniphe Janssens, dorsal view ♂; (172) same, lateral view; (173) L. carotenoides Tomasovic, dorsal view ♀; (174) same, lateral view.

Abbreviations: M3 – third branch of media; myst – mystax; ped – pedicel; pgn – postgena; plp – palpus; pped – postpedicel; pprn – postpronotum; r5 – fifth radial cell; scp – scape; tg – tergite.

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Londt (2010b) and Oldroyd (1970, 1974). Oldroyd (1970, South Africa, Swaziland and Zimbabwe), Central Africa (Ango- 1974) provided the most recent identification keys to part of la, Democratic Republic of Congo, Malawi and Zambia), East the Afrotropical fauna. Africa (Djibouti, Ethiopia, Kenya, Somalia, Sudan, Tanzania and Uganda), the southern Arabian Peninsula (United Arab Irwinomyia Londt (Brachyrhopalinae). An endemic genus Emirates and Yemen; the photograph provided in Fig. 30 also of two small-sized species, I. argentea Londt, 1994 and I. au­ represents a new record for Oman) and West Africa (Burki- rea Londt, 1994, confined to Southern Africa (Namibia) (Londt na Faso, Gambia, Ghana, Mauritania and Niger). The genus 1994b). The biology of the genus is poorly known, but species inhabits savanna and woodland habitats and adults perch at inhabit grassland, savanna and semi-desert habitats and adults the tips of shrubs, bushes and trees (Fig. 30). The oviposition perch on the ground or possibly within and/or at the tips of strategy remains unknown, but probably takes place on or in grass. Oviposition takes place in sand or soil. Londt (1994b) vegetation (wood). Some biological information was provid- provided an identification key to the two species. ed by Bosák & Hradský (2011: 714) and Cuthbertson (1938). Other useful references include Bosák & Hradský (2011), Ischiolobos Londt (Brachyrhopalinae). An endemic genus of Kirk-Spriggs & McGregor (2009), Londt (2010b) and Oldroyd four small-sized species, confined to Southern Africa (Lesotho (1974). Dikow & Londt (2000a) provided an identification key and South Africa) and East Africa (Tanzania) (Londt 2005b). to the four species. The genus inhabits grassland, Fynbos and savanna habitats and adults perch on the ground or within grass. Oviposition takes Laphyctis Loew (Laphriinae). An endemic genus of two place in sand or soil. Other useful references include Lindner medium-sized, wasp-like species (Figs 31, 163, 164), L. gigan­ (1955). Londt (2005b) provided an identification key to the tella (Loew, 1852) and L. orichalcea (Lindner, 1973), requiring four species. modern revision, recorded from Southern Africa (Mozam- bique, Namibia, South Africa and Zimbabwe) and East Africa Juxtasilus Londt (Asilinae). An endemic monotypic genus, (Kenya). The genus apparently inhabits savanna and semi-de- with the single medium-sized species, J. capensis (Londt, sert habitats and adults perch on the ground (Fig. 31). The ovi- 1979), confined to Southern Africa (South Africa) (Londt position strategy remains unknown, but probably takes place 2005a). The species inhabits Fynbos and semi-desert habitats on or in sand or soil. Other useful references include Lindner and adults probably perch within and/or at the tips of shrubs (1973) and Oldroyd (1974), who dealt with species under and bushes. Oviposition takes place on or in vegetation. the name ; later transferred to Laphyctis by Londt (1988b). No identification key is currently available. Katharma Oldroyd (Laphriinae). An endemic monotypic genus, with the single large-sized species, K. sanguinaria Ol- Laphystotes Oldroyd (Laphriinae). An endemic genus of droyd, 1960, confined to the Indian Ocean islands (Mada- two small-sized species (Figs 165, 166, 272–274), L. albicans gascar) (Oldroyd 1960a). The biology of the species remains (Engel, 1932) and L. ariel Londt, 2004, confined to Southern unknown, but it probably inhabits forest habitats and adults Africa (Namibia, South Africa and Zimbabwe) (Londt 2004a). perch within trees. The oviposition strategy is unknown, but The genus inhabits savanna habitats and adults perch on the probably takes place on or in vegetation. Other useful refer- ground. Oviposition probably takes place in sand or soil. Other ences include Tomasovic (2014). useful references include Oldroyd (1974). Londt (2004a) pro- vided an identification key to the two species. Katharmacercus Tomasovic (Laphriinae). An endemic ge- nus of two large-sized species, K. flagellata (Oldroyd, 1960) Lasiocnemus Loew (Leptogastrinae). An endemic genus of and K. matilei (Menier & Tsacas, 2001), confined to the Indian eight small- to large-sized species (Figs 167, 168, 278–280), Ocean islands (Madagascar) (Tomasovic 2014). The biology reviewed by Dikow (2007). The genus is widely distributed of the genus remains unknown, but species probably inhab- throughout sub-Saharan Africa, recorded from Southern Af- it forest habitats and adults perch within shrubs and bushes. rica (Botswana, Namibia, Mozambique, South Africa, Swazi- The oviposition strategy is unknown, but probably takes place land and Zimbabwe), Central Africa (Cameroon, Democratic on or in vegetation. Other useful references include Menier & Republic of Congo, Malawi and Zambia), East Africa (Kenya, Tsacas (2001). No identification key to the species is currently Somalia and Tanzania) and West Africa (Côte d’Ivoire, Nigeria available. and Senegal) (Dikow 2007). There are also unpublished re- cords from Ethiopia. The genus inhabits grassland, savanna and Labarus Londt (Asilinae). An endemic monotypic genus, woodland habitats and adults perch at the tips of grass. Ovipo- with the single small-sized species, L. ignota Londt, 2005 (Figs sition takes place through random egg-dropping. Other useful 237–239, 252, 253), confined to Southern Africa (South Afri- references include Janssens (1952). Dikow (2007) provided an ca) (Londt 2005a). The species inhabits Fynbos habitats and identification key to the eight species. adults probably perch within and/or at the tips of shrubs and bushes. Oviposition takes place on or in vegetation. Laxenecera Macquart (Laphriinae). A large genus of 32 small- to large-sized, bee-like species (Figs 32, 169, 170), re- Lamyra Loew (Laphriinae). A genus of four large- to very quiring modern revision. The genus occurs in the Afrotropical large-sized, wasp-like species (Figs 30, 161, 162, 269–271). and Oriental Regions, with highest diversity in the Afrotrop- The genus is endemic to the Afrotropical Region, although one ics, with 29 species. The genus is widespread throughout sub- species, L. vorax Loew, 1858, extends into Israel and Saudi Saharan Africa, recorded from Southern Africa (Botswana, Mo- Arabia in the Palaearctic Region (Dikow & Londt 2000a). The zambique, Namibia, South Africa and Zimbabwe), Central Af- genus is widespread throughout sub-Saharan Africa, record- rica (Angola, Democratic Republic of Congo, Gabon, Malawi ed from Southern Africa (Botswana, Mozambique, Namibia, and Zambia), East Africa (Burundi, Ethiopia, Kenya, Somalia,

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1150 SURICATA 5 (2017)

Tanzania and Uganda), the southern Arabian Peninsula (re- (1970, 1974), Tomasovic (2008a) and Tomasovic & Constant corded from “Arabia”) and West Africa (Côte d’Ivoire, Gambia, (2013). Oldroyd (1974) provided the most recent identifica- Nigeria, Senegal and Sierra Leone). The genus inhabits grass- tion key to Southern African species. land and savanna habitats and adults perch within and at the tips of grass (Fig. 32). The oviposition strategy is unknown, but Leptogaster Meigen (Leptogastrinae). A very large genus may involve oviposition in or on vegetation. Some biological of 204 small- to medium-sized species (Figs 43, 171–174), information was provided by Cuthbertson (1937, 1938). Oth- occurring in all zoogeographical regions, except Antarctica, er useful references include Londt (2010b, 2012a), Oldroyd with 55 species in the Afrotropics. The genus is in need of

Figs 48.175–180. Habitus photographs of Asilidae: (175) Lobus sp., dorsal view ♂; (176) same, lateral view; (177) Lycoprosopa atrimaculata (Hobby), dorsal view ♂; (178) same, lateral view; (179) Macroetra damara Londt, dorsal view ♂; (180) same, lateral view.

Abbreviations: anatg – anatergite; dc s – dorsocentral seta; myst – mystax; pprn – postpronotum; R4 – upper branch of third branch of radius; sctl – scutellum.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1151 modern revision as it is currently paraphyletic, with the Afro- from Southern Africa (South Africa and Zimbabwe), Central tropical species not representing Leptogaster sensu stricto (T. Africa (Democratic Republic of Congo), East Africa (Burundi, Dikow, unpubl.). Oldroyd (1980: 357) catalogued the fauna Rwanda, Somalia, Sudan, Tanzania and Uganda), the south- and the study of Leptogaster type specimens reveals that many ern Arabian Peninsula (United Arab Emirates and Socotra Is. of these require transfer to existing genera, including Lobus (Yemen)), West Africa (Côte d’Ivoire and Gambia) and the In- and Mesoleptogaster (see below), reducing the total number dian Ocean islands (Madagascar). There are also unpublished of Leptogaster species from 74 to 55 (T. Dikow, unpubl.). The records from Botswana, Eritrea, Ethiopia, Lesotho, Mali, Na- genus is widespread throughout the Afrotropics, recorded mibia and Seychelles. The genus inhabits grassland, savanna

Figs 48.181–186. Habitus photographs of Asilidae: (181) Mesoleptogaster sp., dorsal view ♀; (182) same, lateral view; (183) Michotamia fuscifemorata Joseph & Parui, dorsal view ♂ (non-Afrotropical); (184) same, lateral view; (185) Microstylum sp., dorsal view ♂; (186) same, lateral view.

Abbreviations: anatg – anatergite; dc s – dorsocentral seta; fc – face; M1 – first branch of media; myst – mystax; ped – pedicel; pmtcx memb – postmetacoxal membrane; pped – postpedicel; pprn – postpronotum; scp – scape; styl – stylus; tb – tibia.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1152 SURICATA 5 (2017) and woodland habitats and adults perch within grass (Fig. 43). Afrotropical, Oriental and Palaearctic Regions, with 17 species Oviposition takes place through random egg-dropping. Other in the Afrotropics (Martin 1972). Although seven species were useful references include Bosák & Hradský (2011), Janssens catalogued by Oldroyd (1980: 359), there are numerous addi- (1953, 1954, 1955, 1957), Londt (2010b), Martin (1964) and tional species in the Afrotropics. The study of type specimens Oldroyd (1970). No identification key to Afrotropical fauna is has indicated that 11 species should be assigned to the genus currently available. and unidentified material has also revealed additional unde- scribed species (T. Dikow, unpubl.). The genus is fairly widely Lobus Martin (Leptogastrinae). A genus of 31 small- to distributed, recorded from Central Africa (Democratic Repub- medium-sized species (Figs 44, 175, 176), occurring in the lic of Congo), East Africa (Ethiopia, Eritrea, Kenya, Rwanda and

Figs 48.187–192. Habitus photographs of Asilidae: (187) Microstylum sp., dorsal view ♂; (188) same, lateral view; (189) same, dorsal view ♀; (190) same, lateral view; (191) Nannolaphria nigra Londt, dorsal view ♀; (192) same, lateral view.

Abbreviations: acanth sp – acanthophorite spine; anepst – anepisternum; dc s – dorsocentral seta; fc – face; M1 – first branch of media; myst – mystax; pgn – postgena; pmtcx memb – postmetacoxal membrane; pped – postpedicel; r1 – first radial cell; r5 – fifth radial cell.

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Somalia) and West Africa (Liberia). There are also unpublished random egg-dropping. Other useful references include Tomas- records from Angola, Benin, Cameroon, Central African Re- ovic (2013) and Tomasovic & De Bakker (2010). No identifica- public, Ghana, Guinea-Bissau, Madagascar, Namibia, Nigeria, tion key to Afrotropical species is currently available. South Africa, Tanzania, Uganda and Zambia. Recorded local- ities indicate that the genus inhabits forest or woodland hab- Loewinella Hermann (Laphriinae). A genus of nine small- itats and adults probably perch at the tips of shrubs, bushes sized species, occurring in the Afrotropical and Palaearctic and trees (Fig. 44). Oviposition probably takes place through Regions, with six species in the Afrotropics (Londt 1982a).

Figs 48.193–198. Habitus photographs of Asilidae: (193) Neolophonotus bimaculatus Londt, dorsal view ♂; (194) same, lateral view; (195) N. chionthrix Hull, dorsal view ♂; (196) same, lateral view; (197) Oligopogon penicillatus Loew, dorsal view ♂; (198) same, lateral view.

Abbreviations: al – alula; anatg – anatergite; dc s – dorsocentral seta; fc – face; pocl s – postocular seta; pped – postpedicel; prepst

– proepisternum; prst – prosternum; r5 – fifth radial cell; sct mn – scutal mane; styl – stylus; term – terminalia.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1154 SURICATA 5 (2017)

The genus is widespread in the Afrotropics, recorded from grassland and semi-desert habitats and adults perch on the Southern Africa (Mozambique, Namibia, South Africa and ground. Oviposition takes place in sand or soil. Londt (1994b) Zimbab­we), Central Africa (Democratic Republic of Congo), provided an identification key to the three species. West Africa (Côte d’Ivoire, Niger and Nigeria) and the Indian Ocean islands (Madagascar). The genus inhabits grassland, sa- Megadrillus Bigot (Asilinae). An endemic genus of two vanna and woodland habitats and adults perch at the tips of medium-sized species, M. brevipennis (Macquart, 1838) and grass. Oviposition probably takes place through random egg- M. heteronevrus (Macquart, 1838), transferred to Neolophono­ dropping. Londt (1982a) provided an identification key to the tus and reviewed by Londt (1987b), but subsequently resurrect- six Afrotropical species. ed by Londt (2004b). The genus is confined to Southern Africa (South Africa) and inhabits Fynbos habitats and adults perch on Longibeccus Scarbrough (Ommatiinae). An endemic genus the ground or on stones. Oviposition takes place in sand or soil. of two medium- to large-sized species (Figs 284–286, 290, Species can be identified by reference to Londt (1987b). 291), L. fuscovittatus (Ricardo, 1900) and L. imperator (Old- royd, 1939), widely distributed throughout sub-Saharan Afri- Melouromyia Londt (Asilinae). An endemic genus of two ca, recorded from Southern Africa (Botswana, Mozambique, medium-sized species (Fig. 9), M. Londt, 2002 and South Africa and Zimbabwe), Central Africa (Angola, Chad and M. natalensis (Ricardo, 1919), recorded from Southern Africa Democratic Republic of Congo), East Africa (Ethiopia, Kenya, (Botswana, Mozambique, South Africa, Swaziland and Zimbab- Sudan and Uganda) and West Africa (Côte d’Ivoire, Ghana we) and Central Africa (Malawi) (Londt 2002b). The genus in- and Nigeria) (Scarbrough 2010). The genus inhabits savanna habits forest, savanna and woodland habitats and adults perch and woodland habitats and adults probably perch within grass, within and at the tips of shrubs and bushes. Oviposition proba- shrubs and bushes. Oviposition probably takes place through bly takes place on or in vegetation. Londt (2002b) provided an random egg-dropping. Scarbrough (2010) provided an identi- identification key to the two species. fication key to the two species. Metommatius Hull (Ommatiinae). A genus of three medium- Lycoprosopa Hull (Asilinae). An endemic genus of two sized species, M. planatus (Scarbrough & Marascia, 2000), large-sized species (Figs 177, 178), L. atrimaculata (Hob- M. politus (Scarbrough & Marascia, 2000) and M. pulchellus by, 1934) and L. dioctriaeformis (Macquart, 1846), requiring (Bromley, 1936), occurring in the Afrotropical and Palaearctic modern revision. The genus is confined to the Indian Ocean Regions (Scarbrough & Marascia 2000). The genus is wide- islands (Madagascar and Mauritius). The genus probably in- spread throughout sub-Saharan Africa, recorded from South- habits forest habitats and adults probably perch within and/or ern Africa (Botswana, Namibia, South Africa, Swaziland and at the tips of trees. Oviposition takes place on or in vegetation. Zimbabwe), Central Africa (Malawi), East Africa (Kenya and Other useful references include Hobby (1934), Hull (1962) Tanzania) and West Africa (Ghana). The genus inhabits savan- and Londt (2005a). No identification key to the two species is na and woodland habitats and adults perch at the tips of grass, currently available. shrubs, bushes and trees. Oviposition takes place through ran- dom egg-dropping. Scarbrough (2010) published an identifica- Lycostommyia Oldroyd (Tillobromatinae). An endemic ge- tion key to the three species. nus of six medium-sized species, confined to Southern Africa (Namibia and South Africa) (Londt 1992a). The genus inhabits Mesoleptogaster Frey (Leptogastrinae). A genus of 30 medium- Fynbos, grassland, savanna, semi-desert and woodland habi- sized species (Figs 45, 181, 182), occurring in the Afrotropical, tats and adults perch on the ground or on rocks. Oviposition Oceanian, Oriental and Palaearctic Regions. Five species occur takes place in sand or soil. Other useful references include in the Afrotropics. Hull (1962) accepted Frey’s subgenus and Hermann (1907) and Oldroyd (1974). Londt (1992a) provided combined Leptogaster madagascariensis Frey, 1937 with Mes­ an identification key to the six species. oleptogaster, which was not followed by Oldroyd (1980: 357). The genus was, however, accepted by Geller-Grimm (2004) and Machimus Loew (Asilinae). A genus of 187 medium-sized spe- Lehr (1988a: 270). This species, along with four other Lepto­ cies, occurring in the Afrotropical, Nearctic, Oriental, Palaearc- gaster species and additional undescribed species requires to tic and possibly Neotropical Regions, but requiring modern be combined with Mesoleptogaster, based on the study of type revision. Thirteen species occur in the Afrotropics, confined to specimens of Leptogaster (T. Dikow, unpubl.). Mesoleptogaster is Central Africa (Democratic Republic of Congo and Malawi) and a fairly widely distributed genus, with unpublished records from East Africa (Ethiopia, Kenya, Tanzania (incl. Zanzibar Is.) and Central Africa (Angola, Cameroon and Democratic Republic of Uganda). The genus probably inhabits savanna and woodland Congo), East Africa (Ethiopia and Tanzania), the Indian Ocean habitats and adults perch on the ground (Efflatoun 1934, 1937) islands (Madagascar), Southern Africa (Mozambique and South and within shrubs and bushes. Oviposition takes place on or in Africa) and West Africa (Ghana, Mali and Nigeria). The biology vegetation. Some biological information was provided (as Tol­ remains unknown. Recorded localities suggest that the genus in- merus) by Cuthbertson (1935, 1936). Other useful references habits forest or woodland habitats and adults probably perch at include Londt (2002b) and Tomasovic (2013, 2014). No identi- the tips of shrubs, bushes and trees (Fig. 45). Oviposition proba- fication key to Afrotropical species is currently available. bly takes place through random egg-dropping. No identification key to Afrotropical species is currently available. Macroetra Londt (Brachyrhopalinae). An endemic genus of three small-sized species (Figs 179, 180), M. angola Londt, Michotamia Macquart (Ommatiinae). A genus of 34 large- 1994, M. cera Londt, 1994 and M. damara Londt, 1994, re- sized species (Figs 183, 184), occurring in the Afrotropical, corded from Southern Africa (Namibia and South Africa) and Australasian and Oriental Regions, with the single species, M. Central Africa (Angola) (Londt 1994b). The genus inhabits coarctata (Macquart, 1855), recorded from the Afrotropics

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(Scarbrough 2010), confined to the Indian Ocean islands Microphontes Londt (Brachyrhopalinae). An endemic genus (Madagascar). The biology remains unknown, but the genus of three small-sized species, M. megoura Londt, 1994, M. safra probably inhabits forest habitats with adults perching at the Londt, 1994 and M. whittingtoni Londt, 1994, confined to tips of shrubs, bushes, or trees. Oviposition probably takes Southern Africa (Namibia and South Africa) (Londt 1994b). The place through random egg-dropping. genus inhabits Fynbos and grassland habitats and adults perch

Figs 48.199–204. Habitus photographs of Asilidae: (199) Pegesimallus calvifrons Londt, dorsal view ♂; (200) same, lateral view; (201) Philodicus fraterculus (Walker), dorsal view ♂; (202) same, lateral view; (203) Promachus amastrus (Walker), dorsal view ♂; (204) same, lateral view.

Abbreviations: anatg – anatergite; aux vn – auxiliary vein; C – costal vein; cua – anterior cubital cell; dc s – dorsocentral seta; fc – face;

M1 – first branch of media; M2 – second branch of media; m3 – third medial cell; myst – mystax; oc tr – ocellar triangle; oc tub – ocellar tubercle; pprn lb – postpronotal lobe; r1 – first radial cell; R1+R2+3 – anterior branch of radius + second branch of radius; r2+3 – second + third radial cell; R2+3 – second branch of radius; r4 – fourth radial cell; R4 – upper branch of third branch of radius; r5 – fifth radial cell; R5 – lower branch of third branch of radius; sctl – scutellum; styl – stylus; tb – tibia; tg – tergite; tsm – tarsomere.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1156 SURICATA 5 (2017) on the ground. Oviposition takes place in sand or soil. Londt with many undescribed species known from Madagascar alone (1994b) provided an identification key to the three species. (E. Fisher, pers. comm. 2017). Seventy-nine species occur in the Afrotropics, but 50 valid names are catalogued and species Microstylum Macquart (Stenopogoninae). A very large ge- remain in need of modern taxonomic treatment. The genus is nus of 138 large- to very large-sized species (Figs 53–55, 74, widespread throughout the Afrotropics, recorded from South- 185–190), occurring in the Afrotropical, Nearctic, Neotropi- ern Africa (Lesotho, Mozambique, Namibia, South Africa and cal (E. Fisher, pers. comm. 2017), Oriental and Palaearctic Zimbabwe), Central Africa (Angola, Chad, Democratic Repub- Regions, while Geller-Grimm (2004) questioned records from lic of Congo, Gabon, Malawi and Zambia), East Africa (Ken- the Australasian Region. The genus requires modern revision, ya, Sudan and Tanzania), West Africa (Gambia, Nigeria and

Figs 48.205–210. Habitus photographs of Asilidae: (205) Prytanomyia kochi (Lindner), dorsal view ♀; (206) same, lateral view; (207) Pycnomerinx sp., dorsal view ♂; (208) same, lateral view; (209) Remotomyia longipalpus Londt, dorsal view ♂; (210) same, lateral view.

Abbreviations: m3 – third medial cell; myst – mystax; prbs – proboscis.

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Senegal) and the Indian Ocean islands (Madagascar). The ge- further five new species. Bromley (1942), Engel (1932), Old- nus inhabits forest, Fynbos, grassland, savanna and woodland royd (1960a) and Timon-David (1952) together added a fur- habitats and adults perch on the ground, on rocks, or at the ther nine species to the fauna, Timon-David (1952) producing tips of grass, shrubs or bushes (Figs 53–55). Oviposition takes another identification key to species, which Oldroyd (1960a) place in sand or soil. Some biological information was provid- updated. Malagasy Microstylum are among the largest known ed by Cuthbertson (1933, 1937, 1939). The Malagasy fauna Afrotropical Asilidae and a specimen of Microstylum magnum has received useful taxonomic treatments through the work of Bromley, 1927 in Muséum national d’Histoire naturelle, Par- Bromley (1927), who described seven new species and keyed is, France, has a body length of 65 mm and a wing length of 11 of the 15 then known and Bromley (1931), who added a 40 mm (E. Fisher, pers. comm. 2017). Other useful references

Figs 48.211–216. Habitus photographs of Asilidae: (211) Rhabdogaster sp., dorsal view ♂; (212) same, lateral view; (213) Rhadi­ nus tewfiki Efflatoun, dorsal view ♀; (214) same, lateral view; (215) Rhipidocephala sp., dorsal view ♂; (216) same, lateral view.

Abbreviations: C – costal vein; CuP – posterior branch of cubital vein; cup – posterior cubital cell; m3 – third medial cell; myst – mystax; pmtcx brg – postmetacoxal bridge; pped – postpedicel; pulv – pulvillus; styl – stylus; tg – tergite.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1158 SURICATA 5 (2017) include Engel’s (1932) key to the species he was familiar with, grassland habitats and adults perch within or at the tips of Bromley (1947) and Oldroyd (1970). No single identification grass. Oviposition takes place on or in vegetation, with the eggs key to Afrotropical species is currently available. pasted between leaves (Londt & Harris 1987). Londt (2014a) provided an identification key to the four species. Millenarius Londt (Asilinae). An endemic genus of four medium-sized species (Figs 240–242, 318, 319), described by Nannolaphria Londt (Laphriinae). An endemic monotypic Londt (2005a) and later reviewed by Londt (2014a). The ge- genus, with the single medium-sized species, N. niger Londt, nus is confined to Southern Africa (South Africa) and inhabits 1977 (Figs 191, 192), confined to Southern Africa (South Africa)

Figs 48.217–222. Habitus photographs of Asilidae: (217) Saropogon zinidi Londt, dorsal view ♀; (218) same, lateral view; (219) Scylaticus costalis (Wiedemann), dorsal view ♂; (220) same, lateral view; (221) Sisyrnodytes subater Oldroyd, dorsal view ♀; (222) same, lateral view.

Abbreviations: anatg – anatergite; C – costal vein; CuA+CuP – anterior branch of cubital vein + posterior branch of cubital vein;

cup – posterior cubital cell; fc – face; myst – mystax; pped – postpedicel; r5 – fifth radial cell; sctl – scutellum; styl – stylus; tb – tibia.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1159

(Londt 1977, 2015a). The species inhabits forest habitats and sandy areas in open forest and woodland habitats and adults adults perch at the tips of shrubs and bushes. Oviposition perch on the ground, or on riverbanks and beaches. Oviposi- probably takes place on or in vegetation. tion takes place in sand or soil.

Nanoculcita Londt & Copeland (Stichopogoninae). An en- Neolophonotus Engel (Asilinae). A near endemic genus demic monotypic genus, with the single medium-sized spe- of 256 medium- to large-sized species (Figs 10–12, 75, 193– cies, N. kenya Londt & Copeland, 2017, confined to Eastern 196, 243–245), 254 of which occur in the Afrotropical Re- Africa (Kenya) (Londt & Copeland 2017). The species inhabits gion, with two species extending into Egypt in the Palaearctic.

Figs 48.223–228. Habitus photographs of Asilidae: (223) Sporadothrix gracilis Hermann, dorsal view ♂; (224) same, lateral view; (225) Stichopogon hermanni Bezzi, dorsal view ♂; (226) same, lateral view; (227) Stiphrolamyra diaxantha (Hermann), dorsal view ♂; (228) same, lateral view.

Abbreviations: al – alula; anepst – anepisternum; C – costal vein; CuA+CuP – anterior branch of cubital vein + posterior branch

of cubital vein; cup – posterior cubital cell; fem – femur; M2 – second branch of media; m3 – third medial cell; plp – palpus; prbs – proboscis; R4 – upper branch of third branch of radius; sct – scutum; st vn – stump vein; tb – tibia.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1160 SURICATA 5 (2017)

Neolophonotus is one of the most speciose genera globally and Oligopogon Loew (incertae sedis). A genus of 35 small-sized is divided into five species-groups, based on overall morpho- species (Figs 24, 197, 198), occurring in the Afrotropical and logical similarity. The genus was reviewed by Londt (1985d, Palaearctic Regions, with 29 species in the Afrotropics (Londt 1986b, 1987b, 1988a), who subsequently added a few ad- 2014b). The taxonomic position of the genus remains uncer- ditional species (Londt 1990a) and is widespread throughout tain. Oldroyd (1980: 370) placed it together with genera he sub-Saharan Africa, recorded from Southern Africa (Botswa- listed as Trigonomiminae; Geller-Grimm (2004) listed it under na, Lesotho, Mozambique, Namibia, South Africa, Swaziland the subfamily Stenopogoninae; and Dikow (2009a) considered and Zimbabwe), Central Africa (Angola, Democratic Republic the genus as incertae sedis. The genus has its highest species di- of Congo, Malawi and Zambia), East Africa (Eritrea, Ethiopia, versity in the Afrotropics, where it is widespread throughout the Kenya, Somalia, Tanzania and Uganda) and the southern Ara- region, recorded from Southern Africa (Lesotho, Mozambique, bian Peninsula (United Arab Emirates). The genus inhabits, Namibia, South Africa, Swaziland and Zimbabwe), Central Af- grassland, Fynbos, savanna, semi-desert and woodland habi- rica (Angola, Democratic Republic of Congo, Malawi and Zam- tats and adults perch on the ground, on beaches, rocks and bia), East Africa (Burundi, Kenya, Sudan and Tanzania), West stones, in riparian habitats or at the tips of grass, shrubs, bushes Africa (Côte d’Ivoire, Gambia, Ghana, Nigeria and Senegal) and and trees (Figs 10–12). Oviposition takes place in sand or soil, the Indian Ocean islands (Madagascar) (Londt 2014b). The ge- or on or in vegetation (N. suillus species-group). Some biolog- nus inhabits forest (margins), grassland and savanna habitats and ical information was provided by Cuthbertson (1936, 1937, adults perch on the ground, within or at the tips of grass, shrubs, 1938, 1939, some as Lophopeltis Engel). Other useful refer- bushes and trees (Fig. 24). Oviposition probably takes place in ences include Bosák & Hradský (2011), Engel (1927), Londt sand or soil. Other useful references include Geller-Grimm & (2004b), Tomasovic (2009) and Tomasovic & Constant (2013). Hradský (2003), Londt (2010b, 2012a), Oldroyd (1970, 1974) Londt (1985d, 1986b, 1987b, 1988a, 1990a) provided the and Tomasovic & De Bakker (2010). Londt (2014b) provided most recent identification keys to Afrotropical species, but see an identification key to the 29 Afrotropical species. also Megadrillus (Londt 2004b). Ommatius Wiedemann (Ommatiinae). A very large genus of Notiolaphria Londt (Laphriinae). An endemic genus of six ca 250 medium- to large-sized species (Figs 47, 48, 287–289), medium-sized species (Figs 33, 34), described by Londt (1977) occurring in the Afrotropical, Australasian, Nearctic, Neotropi- and reviewed by Londt (2015a). The genus is recorded from cal and Palaearctic Regions, with 70 species in the Afrotropics. Southern Africa (Mozambique and Zimbabwe), Central Africa The genus requires modern revision. The Malagasy fauna was studied by Oldroyd (1960b) who described four new species (Malawi), East Africa (Burundi, Kenya, Tanzania and Uganda) and later reviewed by Martin (1964), who added a further 20 and the Indian Ocean islands (Comoros, Madagascar, Mauri- species and provided an identification key. The 41 species tius and Réunion Is.) (Londt 2015a). The genus inhabits forest occurring in sub-Saharan Africa still require modern revision, habitats and adults perch on fallen trees (E. Fisher, pers. comm. despite Scarbrough (2002, 2003, 2010) and Scarbrough et al. 2017), but have also been photographed on exposed leaves (2003) having described 19 species in recent years. The ge- (Figs 33, 34). Oviposition probably takes place on or in vege- nus is widespread throughout the Afrotropics, recorded from tation. Londt (2015a) provided an identification key to the six Southern Africa (Mozambique, South Africa and Zimbabwe), species. Central Africa (Angola, Cameroon, Chad, Democratic Repub- Notomochtherus Londt (Asilinae). An endemic monotypic lic of Congo, Malawi, Republic of Congo and Zambia), East Af- rica (Burundi, Ethiopia, Kenya, Rwanda, Sudan, Tanzania and genus, with the single medium-sized species, N. brevicauda Uganda), the southern Arabian Peninsula (United Arab Emir- Londt, 2002, confined to Southern Africa (South Africa) (Londt ates and Yemen (incl. Socotra Is. and Abd al Kuri Is.)), West 2002b). The species probably inhabits Fynbos and grassland Africa (Côte d’Ivoire, Gambia, Ghana, Guinea, Niger, Nigeria habitats and adults perch within grass. The oviposition strategy and Senegal) and the Indian Ocean islands (Comoros, Mada- remains unknown as females are unknown. gascar and Mauritius). The genus inhabits grassland, savanna Nusa Walker (Laphriinae). A genus of 30 medium-sized and woodland habitats and adults perch within grass and at species (Fig. 35), occurring in the Afrotropical, Oriental and the tips of shrubs, bushes and trees (Figs 47, 48). Oviposition Palaearctic Regions, with three species in the Afrotropics, N. eos takes place through random egg-dropping. Some biological in- Londt, 2006, N. infumata (Loew, 1851) and N. ingwavuma Old- formation was provided by Cuthbertson (1933, 1934, 1936, 1937, 1938, 1939). Other useful references include Bromley royd, 1974 (Londt 2006a). The genus is widespread throughout (1936, 1942), Londt (2010b, 2012a), Oldroyd (1960b, 1968, sub-Saharan Africa, recorded from Southern Africa (Botswana, 1970, 1974) and Tomasovic (2009). No single identification Mozambique, Namibia, South Africa and Zimbabwe), Central key to Afrotropical species is currently available. Africa (Democratic Republic of Congo, Malawi and Zambia), East Africa (Ethiopia, Kenya, Somalia, Sudan, Tanzania (incl. Ontomyia Dikow & Londt (Stenopogoninae). An endemic Zanzibar Is.) and Uganda), the southern Arabian Peninsula (So- monotypic genus, with the single medium-sized species, O. cotra Is. (Yemen)) and West Africa (Gambia, Ghana and Mali). ricardoi (Londt, 1985) (Figs 295–297), confined to Southern The genus inhabits grassland and savanna habitats and adults Africa (Namibia) (Dikow & Londt 2000b). The species inhabits perch on the ground and on stones (Fig. 35). Oviposition prob- grassland, savanna and semi-desert habitats and adults perch ably takes place in sand or soil. Some biological information on the ground. Oviposition takes place in sand or soil. was provided by Cuthbertson (1939, under the synonym Da­ sythrix Loew). Other useful references include Londt (2010b) Oratostylum Ricardo (Stenopogoninae). An endemic genus and Oldroyd (1970, 1974). Londt (2006a) provided an identifi- of three medium-sized species, O. crenum Dikow & Londt, cation key to the three Afrotropical species. 2000, O. lepidum Ricardo, 1925 and O. zebra Dikow & Londt,

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1161

2000, confined to Southern Africa (Botswana, Namibia, South Orthogonis Hermann (Laphriinae). A genus of 15 medium- Africa and Zimbabwe) (Dikow & Londt 2000b). The genus in- sized species, occurring in the Afrotropical, Australasian, habits grassland and savanna habitats and adults perch on the Nearctic and Oriental Regions, with the single Afrotropical ground. Oviposition takes place in sand or soil. Other useful species, O. madagascarensis Bromley, 1942, confined to the references include Ricardo (1925). Dikow & Londt (2000b) Indian Ocean islands (Madagascar) (Bromley 1942). The biol- provided an identification key to the three species. ogy remains unknown, but the species probably inhabits forest

Figs 48.229–234. Habitus photographs of Asilidae: (229) Synolcus acrobaptus (Wiedemann), dorsal view ♂; (230) same, lateral view; (231) Torasilus solus Londt, dorsal view ♂; (232) same, lateral view; (233) Trichardis picta Hermann, dorsal view ♂; (234) same, lateral view.

Abbreviations: anatg – anatergite; C – costal vein; cua – anterior cubital cell; CuA+CuP – anterior branch of cubital vein + poste- rior branch of cubital vein; cx – coxa; d – discal cell; dc s – dorsocentral seta; fc – face; fem – femur; pocl s – postocular seta;

pped – postpedicel; pprn lb – postpronotal lobe; R1 – anterior branch of radius; R2+3 – second branch of radius; r5 – fifth radial cell; scp – scape; styl – stylus.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1162 SURICATA 5 (2017) habitats and adults probably perch within or at the tips of on the ground, within grass or within and at the tips of shrubs shrubs, bushes and trees. The oviposition strategy is unknown, and bushes (Figs 21–23). Oviposition takes place in sand or but probably takes place on or in vegetation. Other useful ref- soil. Some biological information was provided by Cuthbertson erences include Hull (1962). (1936, 1938, as Neolaparus Williston). Other useful references include Bromley (1936), Londt (2010b, 2012a) and Tomasovic Oxynoton Janssens (Brachyrhopalinae). An endemic genus (2008b). Londt (1980b) provided an identification key to the of two small- to medium-sized species, O. arnaudi Oldroyd, 46 Afrotropical species. 1974 and O. francoisi Janssens, 1951, recorded from South- ern Africa (Zimbabwe), Central Africa (Chad, Democratic Re- Perasis Hermann (Laphriinae). A genus of ten medium-sized public of Congo and Zambia) and East Africa (Burundi, Kenya, species, occurring in the Afrotropical, Nearctic, Neotropical Rwanda and Tanzania) (Londt 1996). Oldroyd (1980: 371) and Palaearctic Regions. Two species occur in the Afrotrop- listed the genus in the tribe Xenomyzini (Trigonomiminae), but ics, P. carpenteri Oldroyd, 1970 and P. transvaalensis Ricardo, Londt (1996) included it to the subfamily Stenopogoninae. It is 1925 (Londt 2007a). The genus is fairly widespread in sub- here included in the subfamily Brachyrhopalinae. The biology Saharan Africa, recorded from Southern Africa (South Africa) and immature stages remain unknown, but the genus proba- and East Africa (Burundi and Uganda). Lehr (1988a: 211) further bly inhabits savanna habitats and adults probably perch on the records P. arabicus (Macquart, 1838) from the southern Arabi- ground and within and at the tips of grass. Oviposition takes an Peninsula (Yemen), but no specimens from the Afrotropical place in sand or soil. Londt (1996) provided an identification part of the Arabian Peninsula were studied by Londt (1999b, key to the two species. 2007a). The genus inhabits grassland and savanna habitats and adults perch on the ground. Oviposition probably takes place Pedomyia Londt (Brachyrhopalinae). An endemic genus of in sand or soil. Other useful references include Londt (1999b), nine small-sized, bee-like species, confined to Southern Africa Oldroyd (1970) and Ricardo (1925). Londt (2007a) provided (Namibia and South Africa) (Londt 1994b). The genus inhabits an identification key to the two Afrotropical species. grassland, Fynbos, savanna and semi-desert habitats and adults perch on the ground. Oviposition takes place in sand or soil. Philodicus Loew (Asilinae). A genus of 50 large-sized spe- Londt (1994b) provided an identification key to the nine spe- cies (Figs 13, 201, 202, 246–248) requiring modern revision, cies. occurring in the Afrotropical, possibly Australasian, Oriental and Palaearctic Regions, with 23 species in the Afrotropics Pegesimallus Loew (Dasypogoninae). A large genus of 51 (Blasdale 1957). The genus is widespread throughout sub- small- to large-sized species (Figs 21–23, 76, 199, 200, 262, Saharan Africa, recorded from Southern Africa (Botswana, Le- 263), occurring in the Afrotropical, Oriental and Palaearctic sotho, Namibia, South Africa and Zimbabwe), Central Africa Regions, with 46 species in the Afrotropics (Londt 1980b). The (Democratic Republic of Congo and Malawi), East Africa (Ken- genus is widespread, recorded from Southern Africa (Botswa- ya, Somalia, Sudan and Uganda), the southern Arabian Pen- na, Lesotho, Mozambique, Namibia, South Africa, Swaziland insula (Yemen) and West Africa (Benin, Côte d’Ivoire, Gam- and Zimbabwe), Central Africa (Angola, Cameroon, Central bia, Ghana, Guinea, Liberia, Nigeria, Senegal, Sierra Leone African Republic, Democratic Republic of Congo, Gabon, Ma- and Togo). The genus probably inhabits grassland (usually in lawi and Zambia), East Africa (Burundi, Kenya, Somalia, Su- the vicinity of fresh or saline water), savanna and woodland dan, Tanzania and Uganda), West Africa (Benin, Burkina Faso, habitats and adults perch on the ground or on beaches and Côte d’Ivoire, Gambia, Guinea, Nigeria, Senegal and Sierra Le- river banks and at the tips of grass, shrubs and bushes (Fig. one) and the Indian Ocean islands (Madagascar). The genus in- 13). Oviposition takes place in sand or soil. Some biological habits forest, savanna and woodland habitats and adults perch information was provided by Cuthbertson (1937, 1938, 1939).

Figs 48.235–236. Habitus photographs of Asilidae: (235) Trichoura tankwa Londt, dorsal view ♂; (236) same, lateral view.

Abbreviations: cua – anterior cubital cell; m3 – third medial cell; myst – mystax.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1163

Other useful references include Londt (1978, 2005a, 2010b, Proagonistes Loew (Laphriinae). A large endemic genus 2012a, 2015b) and Tomasovic (2012a). Blasdale (1957) and of 22 large- to very large-sized, pompilid-mimicking species Londt (2015b) provided the most recent identification keys to (Figs 36, 37) reviewed by Bromley (1930, 1933), but requiring Afrotropical species. modern revision. The genus is widely distributed throughout sub-Saharan Africa, recorded from Southern Africa (Mozam- Pilophoneus Londt (Laphriinae). An endemic genus of two bique, South Africa and Zimbabwe), Central Africa (Cam- medium-sized, bee-like species, P. analogos Londt, 2014 eroon, Democratic Republic of Congo, Gabon, Malawi and and P. krugeri (Oldroyd, 1974), described by Londt (1988b) Republic of Congo), East Africa (Kenya, Tanzania and Uganda), and later reviewed by Londt (2014d). The genus is recorded West Africa (Côte d’Ivoire, Ghana, Guinea, Nigeria and Sierra Leone), the Indian Ocean islands (Madagascar) and the Atlan- in sub-Saharan Africa from Southern Africa (South Africa), tic Ocean islands (São Tomé and Príncipe). The genus inhabits Central Africa (Zambia) and East Africa (Tanzania). The genus forest and woodland habitats and adults perch within shrubs, inhabits grassland and savanna habitats and adults probably bushes and trees (Figs 36, 37). Other useful references include perch within and/or at the tips of grass. The oviposition strategy Oldroyd (1970), Tomasovic (2010) and Tomasovic & Kwandjo remains unknown, but may involve inserting eggs into dead (2007). Oviposition probably takes place on or in vegetation. wood. Other useful references include Oldroyd (1974). Londt Oldroyd (1970) provided the most recent identification key to (2014d) provided an identification key to the two species. the 22 Afrotropical species.

Figs 48.237–245. Male terminalia of Asilidae (Asilinae): (237) Labarus ignota Londt, lateral view; (238) same, dorsal view; (239) same, ventral view; (240) Millenarius graminosus Londt, lateral view; (241) same, dorsal view; (242) same, ventral view; (243) Neolophonotus boa Londt, lateral view; (244) same, dorsal view; (245) same, ventral view. Figs 237–239 (after Londt 2005a, figs 36–38), Figs 240–242 (after Londt 2005a, figs 44–46), Figs 243–245 (after Londt 1988a, figs 56–58).

Abbreviations: hypd – hypandrium; ph – phallus; st – sternite.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1164 SURICATA 5 (2017)

Promachus Loew (Asilinae). A very large genus of 230 me- 1938, 1939, some as Bactria). Other useful references include dium- to large-sized species (Figs 1, 14–16, 77, 84, 203, 204), Bosák & Hradský (2011), Bromley (1931), Londt (2010b), Ol- occurring in all zoogeographical regions, except Antarctica, but droyd (1974) and Tomasovic (2012b, 2014). No identification requiring modern revision. While Geller-Grimm (2004) placed key to Afrotropical species is currently available. Bactria Meigen in synonymy with Promachus, the taxonomic position of these genera has not been adequately resolved, as Prytanomyia Özdikmen (Laphriinae). An endemic mono- Bactria is the older name (Londt 2005a). Ninety-eight species typic genus, with the single small-sized species, P. kochi (Lind- occur in the Afrotropics, being widespread throughout the ner, 1973) (Figs 38, 78, 205, 206), described by Oldroyd Afrotropics, recorded from Southern Africa (Lesotho, Mozam- (1974, as Prytania). The species is confined to Southern Africa bique, Namibia, South Africa and Zimbabwe), Central Africa (Namibia) and Central Africa (Angola) and inhabits desert hab- (Angola, Bioko Is. (Equatorial Guinea), Cameroon, Democratic itats and adults perch on the ground or on beaches (Fig. 38). Republic of Congo, Gabon, Malawi, Republic of Congo and Oviposition probably takes place in sand or soil. The genus was Zambia), East Africa (Ethiopia, Kenya, Sudan, Tanzania and reviewed by Londt & Dikow (2017). Uganda), the southern Arabian Peninsula (United Arab Emir- ates and Socotra Is. (Yemen)), West Africa (Benin, Côte d’Ivo- Pycnomerinx Hull (Stenopogoninae). An endemic genus ire, Gambia, Ghana, Guinea, Liberia, Nigeria, Senegal, Sierra of three medium-sized, wasp-like species, P. cogani Oldroyd, Leone and Togo) and the Indian Ocean islands (Madagascar). 1974, P. gweta Oldroyd, 1974 and P. rhodesii (Ricardo, 1925) The genus inhabits forest, grassland, Fynbos, savanna, semi- (Figs 207, 208), confined to Southern Africa (Botswana, Na- desert and woodland habitats and adults perch on stones, mibia and Zimbabwe) (Londt 1990c), with unpublished re- within grass or at the tips of shrubs and bushes (Figs 1, 14–16). cords from Mozambique. The genus inhabits grassland and Oviposition takes place on or in vegetation. Some biological savanna habitats and adults perch on the ground or at the tips information was provided by Cuthbertson (1934, 1935, 1937, of grass. Oviposition takes place in sand or soil. Other useful

Figs 48.246–251. Male terminalia of Asilidae (Asilinae): (246) Philodicus tenuipes Loew, lateral view; (247) same, dorsal view; (248) same, ventral view; (249) Synolcus acrobaptus (Wiedemann), lateral view; (250) same, dorsal view; (251) same, ventral view. Figs 246–248 (after Londt 2015b, figs 21–23), Figs 249–251 (after Londt 2012d, figs 14–16).

Abbreviations: epand – epandrium; gonst – gonostylus.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1165 references include Oldroyd (1974). Londt (1990c) provided an Scarbrough & Marascia (2003) provided an identification key identification key to the three species. to the 25 Afrotropical species.

Pygommatius Scarbrough & Marascia (Ommatiinae). A Remotomyia Londt (Stenopogoninae). An endemic genus of genus of 39 medium-sized species, occurring in the Afrotrop- four medium- to large-sized species (Figs 209, 210), confined ical and Oriental Regions, with 25 species in the Afrotropics. to Southern Africa (Botswana, Namibia, South Africa and Zim- Originally described by Scarbrough & Marascia (2003), as a babwe) (Londt 1983c). The genus inhabits grassland, savanna subgenus of Ommatius, the group was elevated to generic and semi-desert habitats and adults perch on the ground. Ovi- rank by Scarbrough & Hill (2005). The genus is widespread, position takes place in sand or soil. Other useful references recorded from Southern Africa (Botswana, Mozambique, Na- include Dikow & Londt (2000b) and Hull (1967). Dikow & mibia, South Africa, Swaziland and Zimbabwe), Central Afri- Londt (2000b) provided an identification key to the four spe- ca (Cameroon, Central African Republic, Chad, Democratic cies. Republic of Congo, Gabon, Malawi, Republic of Congo and Zambia), East Africa (Ethiopia, Kenya, Tanzania and Uganda), Rhabdogaster Loew (Brachyrhopalinae). A near endemic West Africa (Benin, Côte d’Ivoire, Gambia, Ghana, Guinea, genus of 39 small- to medium-sized species (Figs 211, 212, Mali, Nigeria and Sierra Leone) and the Indian Ocean islands 259–261), with R. cinerascens (Wulp, 1899) extending into (Madagascar) (Scarbrough & Marascia 2003). The genus in- Saudi Arabia (Londt 2006c) and Iran (Lehr 1988a: 226) in the habits grassland, savanna and woodland habitats and adults Palaearctic (Londt 2006c). The genus is widespread throughout perch at the tips of grass, shrubs, bushes and trees. Oviposition sub-Saharan Africa, recorded from Southern Africa (Botswana, takes place through random egg-dropping. Other useful refer- Lesotho, Mozambique, Namibia, South Africa, Swaziland and ences include Londt (2010b, 2012a) and Scarbrough (2010). Zimbabwe), Central Africa (Angola, Democratic Republic of

Figs 48.252–255. Female ovipositors of Asilidae (Asilinae): (252) Labarus ignota Londt, lateral view; (253) same, dorsal view; (254) Millenarius graminosus Londt, lateral view; (255) same, dorsal view. Figs 252, 253 (after Londt 2005a, figs 41, 42), Figs 254, 255 (after Londt 2005a, figs 49, 50).

Abbreviation: cerc – cercus.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1166 SURICATA 5 (2017)

Congo and Zambia), East Africa (Eritrea, Kenya, Sudan, Tan- unpublished records from Djibouti. The biology of the genus is zania and Uganda), the southern Arabian Peninsula (Yemen poorly known, but species probably inhabit semi-desert hab- and Oman) and West Africa (Côte d’Ivoire and Gambia) (Londt itats and adults probably perch on the ground or on beaches 2006c). The genus inhabits grassland, Fynbos, savanna and and river banks. Oviposition takes place in sand or soil. Other woodland habitats and adults perch on the ground and at the useful references include Bosák & Hradský (2011), Efflatoun tips of grass. Oviposition takes place in sand or soil. Other use- (1934, 1937) and Geller-Grimm (2002). No identification key ful references include Londt (1993b, 2010b) and Tomasovic & to Afrotropical species is currently available. Constant (2013). Londt (2006c) provided an identification key to the 39 Afrotropical species. Rhipidocephala Hermann (Trigonomiminae). An endemic genus of 26 small-sized species (Figs 64, 65, 215, 216), re- Rhacholaemus Hermann (Stenopogoninae). An endem- viewed by Oldroyd (1966), but requiring modern revision. ic genus of nine small- to medium-sized species confined to Geller-Grimm (2004) and Lehr (1988a: 214) recorded the ge- Southern Africa (Namibia and South Africa) (Londt 1999d). nus as questionable from the Palaearctic parts of the Arabian The genus inhabits grassland, Fynbos, savanna and semi- Peninsula. The genus is widespread, recorded from Southern desert habitats and adults perch on the ground. Oviposition Africa (Botswana, Lesotho, Mozambique, South Africa and takes place in sand or soil. Other useful references include Zimbabwe), Central Africa (Democratic Republic of Congo), Oldroyd (1974). Londt (1999d) provided an identification key East Africa (Burundi, Ethiopia, Kenya, Somalia and Tanzania), to the nine species. the southern Arabian Peninsula (Yemen), West Africa (Gambia, Ghana and Nigeria) and the Indian Ocean islands (Madagascar) Rhadinus Loew (Stichopogoninae). A genus of ten small- (Oldroyd 1966). The genus inhabits grassland, Fynbos, savanna sized, bee-like species (Figs 57, 213, 214), occurring in the and woodland habitats and adults perch at the tips of grass or Afrotropical and Palaearctic Regions, which requires mod- within shrubs and bushes (Figs 64, 65). Oviposition takes place ern revision. Four species occur in the Afrotropics, confined through random egg-dropping. Some biological information to East Africa (Sudan) and the southern Arabian Peninsula was provided by Cuthbertson (1935). Oldroyd (1966) provid- (United Arab Emirates and Yemen (incl. Socotra Is.)), with ed the most recent identification key to the 26 species.

Figs 48.256–261. Male terminalia of Asilidae (Brachyrhopalinae): (256) Habropogon capensis Londt, lateral view; (257) same, dorsal view; (258) same, ventral view; (259) Rhabdogaster eremia Londt, lateral view; (260) same, dorsal view; (261) same, ventral view. Figs 256–258 (after Londt 2000b, figs 28–30), Figs 259–261 (after Londt 2006c, figs 21–23).

Abbreviations: goncx – gonocoxite; gonst – gonostylus; hypd – hypandrium.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1167

Robertomyia Londt (Asilinae). An endemic genus of two in the Afrotropical, Neotropical, Oriental and Palaearctic Re- medium- to large-sized species, R. lavignei Londt, 1990 and gions, with 36 species in the Afrotropics (Londt 1992c). The ge- R. snowi Londt, 2010, described by Londt (1990a) and later nus is distributed mainly in Southern Africa, but is widespread reviewed by Londt (2010b). The genus is recorded from East throughout sub-Saharan Africa, recorded from Southern Africa Africa (Somalia) and West Africa (Gambia), with an unpub- (Botswana, Lesotho, Namibia, South Africa and Zimbabwe), lished record from Kenya. The genus probably inhabits savan- Central Africa (Malawi and Zambia), East Africa (Kenya), West na and woodland habitats and adults probably perch at the Africa (Gambia) and the southern Arabian Peninsula (United tips of shrubs and bushes. Oviposition probably takes place on Arab Emirates) (Londt 1992c). The genus inhabits grassland, or in vegetation. Londt (2010b) provided an identification key Fynbos, savanna, semi-desert and woodland habitats and adults to the two species. perch on the ground or at the tips of shrubs and bushes (Fig. 56). Oviposition takes place in sand or soil. Other useful references Saropogon Loew (Dasypogoninae). A genus of 135 small- include Bosák & Hradský (2011), Engel (1932), Londt (2010b) to medium-sized species (Figs 217, 218), occurring in all zoo­ and Tomasovic & Constant (2013). Londt (1992c) provided an geographical regions, except Antarctica. Ten species occur identification key to the 36 Afrotropical species. in the Afrotropics, confined to East Africa (Eritrea, Ethiopia, Kenya, Sudan and Tanzania), the southern Arabian Peninsu- Sisyrnodytes Loew (Willistonininae). A genus of 18 small- to la (Oman, United Arab Emirates and Yemen) and West Africa medium-sized, bee-like species (Figs 67, 79, 221, 222, 310, (Mali, Mauritania, Niger and Senegal) (Londt 1997). The genus 311), occurring in the Afrotropical and Palaearctic Regions, apparently inhabits grassland, savanna and semi-desert habi- with 16 species in the Afrotropics (Londt 2009a). The genus tats and adults probably perch on the ground or on low vege- is widespread throughout sub-Saharan Africa, recorded from tation. Oviposition takes place in sand or soil. Other useful ref- Southern Africa (Botswana, Mozambique, Namibia, South Af- erences include Bosák & Hradský (2011) and Londt (2010b). rica and Zimbabwe), Central Africa (Malawi and Zambia), East Londt (1997) provided the most recent identification key to Africa (Eritrea, Kenya and Tanzania), the southern Arabian Pen- the 10 sub-Saharan species, but see Bosák & Hradský (2011) insula (United Arab Emirates and Yemen) and West Africa (Be- for the Arabian Peninsula fauna. nin, Gambia, Mauritania, Niger and Nigeria) (Londt 2009a). The genus inhabits grassland, Fynbos, savanna, semi-desert Schildia Aldrich (Leptogastrinae). A genus of ten small-sized and woodland habitats and adults perch on the ground or on species (Figs 281–283), occurring in the Afrotropical, Neotrop- stones and rocks (Fig. 67). Oviposition takes place in sand or ical and Oriental Regions, with a single Afrotropical species, S. soil. Other useful references include Bosák & Hradský (2011) adina Dikow & Bayless, 2009, confined to the Indian Ocean and Oldroyd (1957). Londt (2009a) provided an identification islands (Madagascar) (Dikow & Bayless 2009). While many key to the 16 Afrotropical species. extra­limital species inhabit forest habitats, S. adina purportedly inhabits “xeric shrub lands” and adults probably perch at the Smeryngolaphria Hermann (Laphriinae). A genus of 10 tips of shrubs, bushes and trees. The oviposition strategy prob- small- to medium-sized species, occurring in the Afrotropical ably involves random egg-dropping. Dikow & Bayless (2009) and Oriental Regions. Three species occur in the Afrotropics, provided an identification key to the world fauna. S. bicolorala Tomasovic, 2003, S. pallida Bromley, 1935 and S. bromleyi Londt, 1989, reviewed by Londt (1989b) with further Scylaticus Loew (Stenopogoninae). A genus of 48 small- to revision required. The genus is fairly widespread throughout large-sized, bee- or wasp-like species (Figs 56, 219, 220), occurring the Afrotropics, recorded from Central Africa (Democratic

Figs 48.262–263. Male terminalia of Asilidae (Dasypogoninae): (262) Pegesimallus laticornis (Loew), lateral view; (263) same, ventral view. Figs 262, 263 (after Londt 1980b, figs 53, 54).

Abbreviations: epand – epandrium; goncx – gonocoxite; gonst – gonostylus; hypd – hypandrium.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1168 SURICATA 5 (2017)

Republic of Congo), West Africa (Nigeria) and the Indian most recent identification key to Afrotropical species, but see Ocean islands (Madagascar) (Londt 1989b). The biology of the Tomasovic (2003) for an additional species. genus remains unknown, but species probably inhabit forest habitats and adults probably perch within trees. Oviposition Sphagomyia Londt (Asilinae). An endemic genus of three probably takes place on or in vegetation. Other useful refer- small-sized species, S. botswana Londt, 2002, S. gambia Londt, ences include Tomasovic (2003). Londt (1989b) provided the 2012 and S. kenya Londt, 2002, described by Londt (2002b)

Figs 48.264–274. Male terminalia of Asilidae (Laphriinae): (264) Anypodetus leucothrix Londt, lateral view; (265) same, dorsal view; (266) same, ventral view; (267) Hoplistomerus nobilis Loew, lateral view; (268) same, ventral view; (269) Lamyra gulo (Loew), lateral view; (270) same, dorsal view; (271) same, ventral view; (272) Laphystotes ariel Londt, lateral view; (273) same, dorsal view; (274) same, ventral view. Figs 264–266 (after Londt 2000a, figs 19–21), Figs 267, 268 (after Londt 2007b, figs 21, 23), Figs 269–271 (after Dikow & Londt 2000, figs 4–6), Figs 272–274 (after Londt 2004a, figs 17–19).

Abbreviations: epand – epandrium; goncx – gonocoxite; gonst – gonostylus; hypd – hypandrium.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1169 and reviewed by Londt (2012a). Species are recorded from Stenopogon Loew (Stenopogoninae). A large genus of 170 Southern Africa (Botswana and South Africa), East Africa (Ken- medium-sized species, occurring in the Afrotropical, Nearctic, ya) and West Africa (Gambia). The genus inhabits savanna and Oriental and Palaearctic Regions, that is in need of modern woodland habitats and adults probably perch within shrubs revision. Only two species occur in the Afrotropics, S. coxalis and bushes. Oviposition probably takes place on or in vegeta- (Becker, 1922) and S. lehri Londt, 1999, that were reviewed tion. Londt (2012a) provided an identification key to the three by Londt (1999d, 2002a) and are confined to East Africa (Ken- species. ya, Somalia and Sudan). The biology of the genus remains un- known, but species probably inhabit grassland, savanna and Sporadothrix Hermann (Willistonininae). An endem- semi-desert habitats and adults perch on the ground or on low ic monotypic genus, with the single medium-sized species, vegetation. Oviposition takes place in sand or soil. No identifi- S. gracilis Hermann, 1907 (Figs 80, 223, 224), confined to cation key to Afrotropical species is currently available. Southern Africa (Botswana, Namibia and South Africa) (Londt 2010c). The species inhabits grassland, savanna and semi- Stichopogon Loew (Stichopogoninae). A large genus of desert habitats and adults perch on the ground or within grass. 102 small-sized species (Figs 58–60, 81, 225, 226, 301, 302), Oviposition takes place in sand or soil. occurring in all zoogeographical regions, except Antarctica.

Figs 48.275–283. Male terminalia of Asilidae (Leptogastrinae): (275) Euscelidia ochricornis (Loew), lateral view; (276) same, dor- sal view; (277) same, ventral view; (278) Lasiocnemus londti Dikow, lateral view; (279) same, dorsal view; (280) same, ventral view; (281) Schildia adina Dikow & Bayless, lateral view; (282) same, dorsal view; (283) same, ventral view. Figs 275–277 (after Dikow 2003, fig. 23), Figs 278–280 (after Dikow 2007, figs 5–7), Figs 281–283 (after Dikow & Bayless 2009, figs 13–15).

Abbreviations: epand – epandrium; goncx+hypd cplx – gonocoxal + hypandrial complex; hypd – hypandrium; sur – surstylus.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1170 SURICATA 5 (2017)

Seventeen species occur in the Afrotropics (Londt 1979b), that and adults perch on the ground, or on beaches and river banks are widespread throughout the region, recorded from South- (Figs 58–60). Oviposition takes place in sand or soil. Some ern Africa (Botswana, Lesotho, Mozambique, Namibia, South biological information was provided by Cuthbertson (1938). Africa and Zimbabwe), Central Africa (Angola and Democrat- Other useful references include Bosák & Hradský (2011), Effla- ic Republic of Congo), East Africa (Ethiopia and Sudan), the toun (1934, 1937), Howarth (2006), Londt (2010b), Oldroyd southern Arabian Peninsula (United Arab Emirates and Yemen (1970, 1974) and Séguy (1955). Londt (1979b) provided the (incl. Socotra Is.)), West Africa (Gambia, Mauritania and Ni- most recent identification key to eight Afrotropical species, but ger) and the Indian Ocean islands (Madagascar). The genus see the recent addition of the southern Arabian Peninsula fau- inhabits grassland, Fynbos, savanna and semi-desert habitats na by Bosák & Hradský (2011: 746).

Figs 48.284–291. Male and female terminalia of Asilidae (Ommatiinae): (284) male terminalia of Longibeccus fuscovittatus (Ricar- do), lateral view; (285) same, dorsal view; (286) same, ventral view; (287) same, Ommatius neotenellus Bromley, lateral view; (288) same, dorsal view; (289) same, ventral view; (290) female terminalia of L. fuscovittatus, dorsal view; (291) same, ventral view. Figs 284–291 (after Scarbrough 2010, figs 31–33, 74–76, 37, 38, respectively).

Abbreviations: cerc – cercus; epand – epandrium; goncx – gonocoxite; gonst – gonostylus; hypd – hypandrium; st – sternite; tg – tergite.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1171

Stiphrolamyra Engel (Laphriinae). A genus of 15 small- to (1974) and Theodor (1980). Londt (1983a) provided the large-sized, wasp-like species (Figs 39, 82, 227, 228), occur- most recent identification key to the eight Southern African ring in the Afrotropical and Palaearctic Regions. Ten species species. occur in the Afrotropics (Londt 1983a), that are widespread throughout sub-Saharan Africa, recorded from Southern Afri- Storthyngomerus Hermann (Laphriinae). An endemic ca (Botswana, Mozambique, Namibia, South Africa and Zim- genus of five wasp-like species, distributed throughout the babwe), Central Africa (Democratic Republic of Congo), East Afrotropics, recorded from Southern Africa (Botswana, Mo- Africa (Kenya and Uganda) and the southern Arabian Penin- zambique, South Africa and Zimbabwe), Central Africa (Cam- sula (United Arab Emirates and Yemen). The genus inhabits eroon, Democratic Republic of Congo, Gabon and Malawi), arid biomes, including grassland, savanna and semi-desert East Africa (Kenya, Tanzania and Uganda), West Africa (Côte habitats and adults perch on rocks or at the tips of shrubs and d’Ivoire, Gambia, Ghana, Guinea, Liberia, Nigeria and Sier- bushes (Fig. 39). Oviposition probably takes place through ra Leone) and the Indian Ocean islands (Madagascar) (Londt random egg-dropping. Other useful references include Bosák 1998b). The genus inhabits forest habitats and adults perch at & Hradský (2011), Kirk-Spriggs & McGregor (2009), Oldroyd the tips of shrubs, bushes and trees. Oviposition probably takes

Figs 48.292–300. Male and female terminalia of Asilidae (Stenopogoninae): (292) male terminalia of Ancylorhynchus cruciger (Loew), lateral view; (293) same, dorsal view; (294) same, ventral view; (295) same, Ontomyia ricardoi (Londt), lateral view; (296) same, dorsal view; (297) same, ventral view; (298) female terminalia of A. cruciger, lateral view; (299) same, ventral view; (300) same, dorsal view. Figs 292–294, 298–300 (after Londt 2011, figs 61–66), Figs 295–297 (after Dikow & Londt 2000, figs 2–4).

Abbreviations: epand – epandrium; hypd – hypandrium.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1172 SURICATA 5 (2017) place on or in vegetation. Other useful references include Thallosia Oldroyd (Ommatiinae). An endemic monotypic Londt (2010b) and Oldroyd (1970, 1974). Londt (1998b) pro- genus, with the single medium-sized species, T. congoicola Old- vided an identification key to the five species. royd, 1970, confined to Central Africa (Democratic Republic of Congo) and West Africa (Gambia). The species inhabits savanna Synolcus Loew (Asilinae). An endemic genus of 13 medi- and woodland habitats and adults probably perch within grass um- to large-sized species (Figs 17, 229, 230, 249–251), re- or at the tips of shrubs, bushes and trees. Oviposition probably viewed by Londt (1980a, 1990b, 2012d) and recorded from takes place through random egg-dropping. Other useful refer- Southern Africa (Namibia and South Africa) and Central Africa ences include Londt (2010b) and Scarbrough (2010). (Malawi). The genus inhabits forest, savanna, semi-desert and woodland habitats and adults perch on the ground or within Tolmerus Loew (Asilinae). A genus of 73 medium- to large- shrubs, bushes and trees (Fig. 17). Oviposition takes place on sized species, occurring in the Afrotropical, Nearctic and or in vegetation. Londt (2012d) provided an identification key Palaearctic Regions and requiring modern revision. Three spe- to the thirteen species. cies occur in the Afrotropics, T. socotrae Geller-Grimm, 2002, T. unicus (Becker, 1910) and T. wraniki Geller-Grimm, 2002, Systropalpus Hull (Laphriinae). An endemic monotypic confined to the southern Arabian Peninsula (Yemen (incl. So- genus, with the single large-sized species, S. aurivulpes Hull, cotra Is.)). The status of the genus requires clarification. Old- 1962, confined to East Africa (Ethiopia). The biology remains royd (1980: 339) listed Tolmerus as a synonym of Machimus, unknown, but the species probably inhabits forest or wood- but Geller-Grimm (2002) recorded three species of Tolmer­ land habitats and adults probably perch within trees. Oviposi- us from Socotra Is. (Yemen) and Geller-Grimm (2004) listed tion probably takes place on or in vegetation. Tolmerus as a distinct genus. The genus probably inhabits sa- vanna habitats and adults probably perch within shrubs and Teratopomyia Oldroyd (Brachyrhopalinae). An endemic bushes. Oviposition probably takes place on or in vegetation. monotypic genus, with the single medium-sized, wasp-like No identification key to Afrotropical species is currently avail- species, T. cyaneus (F., 1781), confined to Southern Africa (Le- able, but detailed descriptions are provided by Geller-Grimm sotho and South Africa) (Londt 2009b). The species inhabits (2002). Fynbos and savanna habitats and adults perch on the ground or at the tips of grass, shrubs and bushes. Oviposition takes Torasilus Londt (Asilinae). An endemic monotypic genus, place in sand or soil. with the single large-sized species, T. solus Londt, 2005 (Figs

Figs 48.301–306. Male and female terminalia of Asilidae (Stichopogoninae and Tillobromatinae): (301) male terminalia of Stichopogon punctum Loew, lateral view; (302) same, female terminalia, lateral view; (303) male terminalia of Hypenetes stig­ matias Loew, lateral view; (304) same, male hypandrium, ventral view; (305) same, dorsal view; (306) female terminalia of H. stigmatias, dorsal view. Figs 301, 302 (after Londt 1979b, figs 27, 35), Figs 303–305 (after Londt 1985b, figs 77–79), Fig. 306 (after Dikow 2009a, fig. 87).

Abbreviations: acanth sp – acanthophorite spine; apod – apodeme; cerc – cercus; tg – tergite.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1173

231, 232), confined to Southern Africa (Namibia). The species Trichoura Londt (Willistonininae). An endemic genus of sev- inhabits desert and semi-desert habitats and adults perch on en small-sized species (Figs 68, 235, 236), confined to South- the ground. Oviposition probably takes place on or in vege- ern Africa (South Africa). The genus was described by Londt tation. (1994b) and reviewed by Londt & Dikow (2016). Species in- habit grassland, savanna and semi-desert habitats and adults Trichardis Hermann (Laphriinae). A genus of 28 small- to perch on the ground or on low vegetation (Fig. 68). Oviposi- medium-sized, bee-like species (Figs 40, 41, 233, 234), occur- tion takes place in sand or soil. Londt & Dikow (2016) provid- ring in the Afrotropical and Palaearctic Regions. Twenty-five ed an identification key to the seven species. species occur in the Afrotropics (Londt 2008b), that are wide- spread throughout sub-Saharan Africa, reported from Southern Tsacasiella Lehr (Asilinae). An endemic genus of eight me- Africa (Botswana, Lesotho, Mozambique, Namibia, South Afri- dium- to large-sized species reviewed by Tsacas (1969), who ca and Zimbabwe), Central Africa (Chad, Democratic Republic described most of the species in and Londt of Congo and Malawi), East Africa (Eritrea, Ethiopia, Kenya, So- (2002b). The genus is recorded from Southern Africa (Botswa- malia and Tanzania), the southern Arabian Peninsula (United na, Namibia and Zimbabwe), Central Africa (Democratic Re- Arab Emirates and Yemen (incl. Socotra Is. and Abd al Kuri Is.)) public of Congo, Malawi and Zambia) and East Africa (Kenya and West Africa (Burkina Faso, Côte d’Ivoire, Gambia, Mali, and Rwanda). The genus inhabits forest, savanna and wood- Niger, Nigeria and Senegal). There are also unpublished re- land habitats and adults perch within grass, shrubs and bushes. cords from Djibouti. The genus inhabits grassland, Fynbos, sa- Oviposition probably takes place on or in vegetation. Londt vanna, semi-desert and woodland habitats and adults perch on (2002b) provided an identification key to the eight species. the ground and on stones. Oviposition takes place in sand or soil. Other useful references include Bosák & Hradský (2011), Tuberconspicus Tomasovic (Asilinae). An endemic mono- Bosák et al. (2014), Geller-Grimm (2002), Londt (2010b) and typic genus, with the single medium-sized species, T. aethi­ Oldroyd (1974). Londt (2008b) provided an identification key opicus Tomasovic, 2014, confined to East Africa (Ethiopia) to 25 Afrotropical species. (Tomasovic 2014). The species is morphologically similar to

Figs 48.307–311. Male terminalia of Asilidae (Trigonomiminae and Willistonininae): (307) Damalis neavei Londt, lateral view; (308) same, dorsal view; (309) same, ventral view; (310) Sisyrnodytes apicalis Oldroyd, lateral view; (311) same, ventral view. Figs 307–309 (after Londt 1989a, figs 118–120), Figs 310, 311 (after Londt 2009a, figs 8, 9).

Abbreviations: epand – epandrium; goncx – gonocoxite; gonst – gonostylus; hypd – hypandrium; ph – phallus; prct – proctiger.

ASILIDAE (ASSASSIN FLIES OR ROBBER FLIES) 48 1174 SURICATA 5 (2017)

Figs 48.312–319. Larval features of Asilidae: (312) larval habitus of Asilus crabroniformis L., dorsal view (non-Afrotropical); (313) same, Laphria gibbosa L., dorsal view (non-Afrotropical); (314) head capsule and rod of A. crabroniformis, dorsal view; (315) same, L. gibbosa, dorsal view; (316) same, head capsule, lateral view; (317) same, ventral view; (318) head capsule of Millenarius dichaetus Hull, dorsal view; (319) same, ventral view. Figs 312–317 (after Melin 1923, figs 171, 160, 172, 131–133), Figs 318, 319 (after Londt & Harris 1987, figs 6, 7).

Abbreviations: ant – antenna; cr – cranium; lab – labium; lbr – labrum; md – mandible; mx – maxilla; mx plp – maxillary palpus; p spr – posterior spiracle; sg – segment; v plt – ventral plate.

MANUAL OF AFROTROPICAL DIPTERA – VOLUME 2 SURICATA 5 (2017) 1175 species of the diverse genus Promachus. The habitat and ecol- on or in vegetation. Londt (2002b) provided an identification ogy of the species remain unknown. key to the four species.

Turkmenomyia Paramonov (Stichopogoninae). A genus Wadipogon Bosák & Hradský (Brachyrhopalinae). A genus of four small-sized species, occurring in the Afrotropical and of three small-sized species, occurring in the Afrotropical and Palaearctic Regions. A single species, T. paramonovi Bosák Palaearctic Regions. The genus is primarily distributed in Egypt, & Hradský, 2011, occurs in the Afrotropics, confined to the with two species in the Afrotropics, W. pulchrum (Efflatoun, southern Arabian Peninsula (United Arab Emirates) (Bosák & 1937) and W. szpilai Bosák & Hradský, 2011, confined to the Hradský 2011: 751). Turkmenomyia has been treated as a syn- southern Arabian Peninsula (United Arab Emirates) (Bosák & onym of Eremodromus Zimin (Geller-Grimm 2004; Lehr 1979) Hradský 2011: 739). The biology remains unknown, but the and was reinstated to full generic status for Palaearctic species genus probably inhabits semi-desert habitats and adults prob- (Bosák & Hradský 2011: 751). The biology of the species re- ably perch on the ground. Oviposition takes place in sand mains unknown, but is probably similar to Stichopogon, inhab- or soil. Other useful references include Efflatoun (1937). No iting semi-desert habitats and adults perching on the ground. identification key to Afrotropical species is currently available. Oviposition takes place in sand or soil. Zelamyia Londt (Asilinae). An endemic monotypic genus, Valiraptor Londt (Asilinae). An endemic genus of four medi- with the single small-sized species, Z. alyctus Londt, 2005, um- to large-sized species (Londt 2002b), confined to Southern confined to Southern Africa (South Africa) (Londt 2005a). The Africa (Namibia and South Africa). The genus inhabits forest, genus inhabits Fynbos habitats and adults probably perch with- grassland, savanna and woodland habitats and adults probably in and/or at the tips of shrubs and bushes. Oviposition proba- perch within grass, shrubs and bushes. Oviposition takes place bly takes place on or in vegetation.

Figs 48.320–323. Pupal features of Asilidae: (320) pupal habitus of Andrenosoma cruentum (McAtee), dorsal view (non-Afro- tropical); (321) same, lateral view; (322) same, ventral view; (323) same, detail of anterior section, ventral view. Figs 320–323 (photographs © J. Barnes).

Abbreviations: ant proc – antennal process; ap proc – apical process.

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